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Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS l 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR ( 

IVAN MACKENZIE LAMB 


VOLUME 60 


1958 


The New England Botanical Club, Ine. 


8 and 10 West King St., Lancaster. Pa. 
Botanical Museum, Oxford St., Cambridge 38. Mass. 


73 


Rv) 


Hovora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS l 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR, ( 

IVAN MACKENZIE LAMB 


Vol. 60 January, 1958 No. 709 
CONTENTS: 
A New Species of Samolus (Primulaceae) from Mexico. 
Tee B. Channel d oom cc os DD ee ene ic ee 1 
A Cytotaxonomic Study of the Genus Achillea in Pennsylvania. 
Ehgood B. Ehrlé ...... 4 a Ve e BERN renun T 
Species New to the Flora of North or South Carolina. Harry E. 
Ahles, C. Ritchie Bell and Albert E. Radford................. 10 
Bartholomew’s Cobble. R. C. Rollins (Review) ................ 32 


The New England Botanical Club, Ine. 


8 and 10 West King St., Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


RHODORA.—A monthly journal of botany, devoted primarily to the 
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Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 January, 1958 No. 709 


A NEW SPECIES OF SAMOLUS (PRIMULACEAE) 
FROM MEXICO 


R. B. CHANNELL! 


The last comprehensive treatment of the genus Samolus L. 
(anomalous among the Primulaceae by the half-inferior position 
of the ovary) appeared in the monograph of the Primulaceae by 
Pax and Knuth (1905). According to this work only two species 
are characterized by ebracteate pedicels and by the absence of 
staminodia: S. ebracteatus HBK. (1818) [including S. alyssoides 
Heller (1895) and S. cuneatus Small (1897)] and S. cinerascens 
(Robinson) Pax & R. Knuth (1. ¢.; cf. Robinson, 1892). The 
presence of a group of small glandular hairs at the base of each 
corolla-lobe, rendering the corolla-throat more or less pubescent, 
is another feature common to both and unknown elsewhere in 
the genus. No other species of the genus so characterized has 
subsequently been described. The species here proposed as new 
is evidently related to the two above, inasmuch as it shares with 
them the characters thus far enumerated. One of these charac- 
ters has served as the primary basis for establishing a separate 
genus. Thus, while specific affinities are clear, the assignment of 
the new species to genus deserves further comment. 

Publication of the segregate genus Samodia, originally pro- 
posed but not validly published by Baudo (1843), was validated 
by Small (1933), who accorded generic significance almost solely 
to the absence of staminodia. "Typified by Samolus ebracteatus 
HBK. and to date nomenclaturally monotypic, this genus has 
never enjoyed general acceptance. This is partly due to the fact 
that elsewhere in the Primulaceae conservative taxonomists have 


1 Gray Herbarium and Arnold Arboretum. Present address: Department of Biology, 
Vanderbilt University. 


2 Rhodora [ Vor. 60 


not attributed generic importance to the presence of staminodia 
in an otherwise non-staminodial group (cf. Lysimachia [Tourn. ] 
L. subgenus Selencia Bigelow; Steironema Raf.). Asa character, 
moreover, the presence or absence of staminodia in Samolus cuts 
across a long-established line, perhaps demarcating groups equal- 
ly as natural, between those species of herbaceous habit with 
smooth herbage, as exemplified by Samolus ebracteatus, and 
those of suffruticose habit with glandular-porous herbage, as 
exemplified by Samolus cinerascens. Generic segregation on the 
basis of differences in habit or general appearance alone, however, 
seems wholly unjustified. The other characters distinguishing 
Samodia, relative length of tube to lobes of the corolla and level 
of adnation of the stamens, are generally disregarded as repre- 
senting merely matters of degree. Under the circumstances the 
single Linnaean genus Samolus is here recognized. 

'The new species is based upon two unicate collections in the 
Gray Herbarium found among undetermined Samolus specimens 
examined in a survey of the morphologieal variation and geo- 
graphical distribution of S. ebracteatus for the southeastern Unit- 
ed States. These specimens were collected in western Coahuila, 
Mexico, by Reynaldo Santos, under the direction of Robert M. 
Stewart, and bear the latter’s collection numbers: Stewart 2748, 
25 September 1942, holotype—‘Rancho del Coyote (1 km. S.E.) 
gypsum slopes, scarce, ascending, 2 dm., fl. white. Base of moun- 
tains along the eastern margin of the Valle de Acatita.” Stewart 
2805, 2 October 1942, paratype—‘Canon del Agua Grande by 
water, common, fl. white. Canyons in the Sierra del Sobaco a few 
kilometers west of Las Delicias." 


Samolus dichondrifolius Channell, sp. nov... Herbae perennes, caule- 
scentes (laeves, non glanduloso-porosae), 1-3 dm. altae, eaules repentes 
vel decumbentes, apices foliati adscendentes, axillae foliorum superiorum 
racemos graciles nudos gerentes. Caulis glaber, 2.5 dm. longus, vel ultra, 
internodi 4-8 mm. longi, pars vestustior basibus petiolaribus foliorum 
disruptorum vestita, apices fola numerosa alterna vel approximata 
gerentes. Folia (2-)4-7(-9) em. longa, simplicia, glabra, supra griseo- 
viridia, subtus pallida vel glauca, laminae integrae reniformes apiculato- 
mueronatae, 1-3 em. latae, 1-2(-3) cm. longae, basi in petiolis gracilibus 
linearibus complanatis (1-)2-5(-6) cm. longis abrupte contractae. 
Racemi, pedicelli calvcesque minute glanduloso-pubescentes. Racemi 


2? The Latin description was kindly prepared by Dr. R. C. Foster, Gray Herbarium. 


1958] Channell,—New Species of Samolus 3 


SAMOLUS DICHONDRIFOLIUS: Fig. 1, habit (74 natural size); fig. 2, leaf (natural size); fig. 
3, flower (X 5); fig. 4, opened corolla showing stamens and tufts of glandular hairs (X 5); 
fig. 5, flower with corolla removed to show pistil (X 5); fig. 6, fruit after dehiscence (X 5); 
fig. 7, seed (X 20). All drawings made from holotype: Stewart 2748, Gu. 


simplices, 0.5-2.5 dm. longi, pedicelli graciles ebraeteati 5-10 mm. longi, 
divergenti-adscendentes. Calyx herbaceus, campanulatus, lobi triangu- 
lares vel triangulato-lanceolati, acuti, tubum aequantes. Corolla alba 
(vel pallide rubra in sicco), 3-4 mm. diametro, lobi late ovati vel 
oblongo-ovati, obtusi, eroso-dentieulati vel retusi, tubum paene aequ- 
antes, faux paullo constricta interne pubescens. Stamina inclusa, in 
corollae tubum paene ad medium inserta, antherae ovatae, 0.8-0.5 mm. 
longae, filamenta brevia aequantes; staminodia nulla. Ovarium semi- 
inferius, stylus ea. 0.5 mm. longus, stigma apicale, discoideo-capitatum. 


4 Rhodora [ Vor. 60 


Capsula semi-inferior, ca. 3 mm. diametro, muris crassis, in valvis 
triangularibus paullo recurvatis dehiscens. Semina numerosa, atro- 
brunnea vel nigra, complanata, ovoideo-angularia, axis longus 0.25- 
0.5 mm., integumenta polygonali-reticulata. 

Caulescent perennial herbs (leaves smooth, not glandular-porous) 1—3 
dm. in height, with repent or decumbent stems, the leafy tips ascending, 
bearing slender naked racemes from the upper leaf-axils. Stem glabrous, 
2.5 dm. or more in length, the internodes 4-8 mm. long, the older portion 
bearing the petiolar bases of old or broken foliage, the tips bearing 
numerous alternate or approximate leaves. Leaves (2-)4-7(-9) em. 
long, simple, glabrous, gray-green above, pale or glaucous below, the 
reniform entire blades apiculate-mucronate, 1-3 em. wide, 1-2(-3) 
em. long, abruptly contracted into slender, linear, flattened petioles 
(1-)2-5(—6) em. long. Racemes, pedicels and calyces minutely glandular- 
pubescent. Racemes simple, 0.5-2.5 dm. long, the slender ebracteate 
pedicels 5-10 mm. long, spreading-ascending. Calyx herbaceous, cam- 
panulate, the lobes triangular or triangular-lanceolate, acute, equalling 
the tube. Corolla white (or when dry flesh-pink), 3-4 mm. in diameter, 
the lobes broadly ovate or oblong-obovate, obtuse, erose-dentieulate or 
retuse, nearly equalling the tube, the slightly constrieted throat pubes- 
cent within. Stamens included, inserted on the corolla-tube near the 
middle, the anthers ovate, 0.3-0.5 mm. long, equalling the short fila- 
ments; staminodia wanting. Ovary 14 inferior, the style about 0.5 mm. 
long, capped by the discoid-capitate stigma. Capsule 1% inferior, about 
3 mm. in diameter, the free portion thick-walled and dehiscent by 
slightly reeurving triangular valves. Seeds numerous, dark brown or 
black, flattened, ovoid-angular in outline, the long axis 0.25-05 mm., 
the testa polvgonal-retieulate. (Figs. 1-7.) 


Samolus dichondrifolius is more closely related to the wide- 
spread S. ebracteatus than to S. cinerascens. Known only from 
San Luis Potosí, Mexico, where it grows on alkaline plains, the 
latter species is distinguished by the suffruticose, subligneous 
habit, the conspieuously glandular-porous, linear or linear-lance- 
olate, acute leaves, the linear-lanceolate sepals, exceeding in 
length the valves of the mature capsule, by the entire corolla- 
lobes, anthers 1.5 mm. long, and by the non-dilated stigmas. 

Samolus ebracteatus (sensu lato) 1s distributed in Cuba, Flor- 
ida, Texas, Oklahoma, New Mexico and from Coahuila, Nuevo 
León and Tamaulipas southward into Oaxaca, as well as in Baja 
California, Mexico. Considerably variable in habit, this species 
apparently grows equally well in calcareous clay, sandy soils, 
sandy loam, humus and on almost bare, eroded limestone (cf. 
Small, 1934). According to herbarium data the habitats include 


1958] Channell,——New Species of Samolus 5 


pine woods, brackish flats and salt marshes, gypsum ravines, chalk 
bluffs, limestone rocks, edges of streams, ponds and springs, cal- 
careous clay and granitic soils. It is not surprising, therefore, that 
practieal difficulties have arisen in the interpretation of local vari- 
ation, even though the species is typically riparian. This species 
is characterized by the herbaceous habit, glabrous herbage, broad 
spatulate to spatulate-rotund leaves with broad, gradually con- 
tracted, vaginate bases, often decurrent on the stem, the tri- 
angular-ovate sepals, merely equalling the valves of the mature 
capsule, by the erose-denticulate corolla-lobes, anthers only about 
0.75—1.0 mm. long, and by the capitate stigmas. 

The new species is distinguished from all other members of the 
genus by the distinctive leaves. The reniform-apiculate blades 
and the long, linear petioles give the foliage a striking resem- 
blance to that of species of Dichondra (Convolvulaceae) , whence 
the epithet. Although sharing the same fundamental floral struc- 
ture, including the irregularly erose-denticulate corolla-lobes and 
capitate stigmas, S. dichrondrifolius may be distinguished from 
S. ebracteatus by the smaller size of the flowers and by the nar- 
rower, more slender inflorescences, as well as by obvious foliage 
differences. Samolus ebracteatus has corollas 4-9 mm. in diam- 
eter and pedicels 10-20 mm. long; S. dichondrifolius has corollas 
only 3-4 mm. or less in diameter and pedicels only 5—10 mm. long. 
In addition, the corollas of the latter species are white; those of 
S. ebracteatus are generally pigmented, varying in color from 
pink to purple or even red. The anthers of S. dichondrifolius are 
only about 0.3-0.5 mm. long. 

Little is known, except in general terms, about the type locality 
and associates of the new species. The descriptive account by 
Muller (1947) of the vegetation and climate of Coahuila indi- 
cates that the plains and basins of the southern, western and 
northern three-fourths of the state are extremely arid and are 
occupied chiefly by a strictly desert vegetational type designated 
as Chihuahuan Desert Shrub. The vegetation is polymorphie, 
consisting of low, sparse perennials and ephemeral annuals. 
Larrea tridentata (DC.) Cov. is the most characteristic species 
and the variant vegetational types are described by Muller in 
terms of associates of that species and those which occasionally 
replace it. The only known stations for Samolus dichondrifolius 


6 Rhodora [Vor. 60 


are located within this vegetational type in western Coahuila, 
near the southeastern corner of Chihuahua. As already indi- 
cated, the holotype was collected from a gypsum slope; the 
paratype near water. 

The studies of Johnston (1941) of extensive gypsum flats and 
gypseous ridges in the same vicinity have yielded a rather exten- 
sive list of gypsophiles, including species of Dicranocarpus, Sart- 
wellia, Nerisyrenia, Drymaria, Nama, ete. Whether or not 
Samolus dichondrifolius may be classified as a strictly gypso- 
philous species, however, remains to be determined. 

It is to be noted that Samolus ebracteatus also occurs in this 
area, near water, in relatively close proximity to the type locality 
of the new species (cf. Stewart 2729, 2735 and 2942: aH) and 
elsewhere in gypsum ravines (cf. Johnston 3631 and Waterfall 
6142: aH). According to the collection data the two species flower 
simultaneously, at least in western Coahuila. Insofar as present 
collections indicate, however, the two species retain their morpho- 
logical identity and are probably isolated genetically. 


LITERATURE CITED 


Bavpo, F. 1843. Anagalleidarum index. Ann. Sci. Nat. II. 20: 350. 

HELLER, A. A. 1895. Botanical explorations in southern Texas during the 
season of 1894. Contrib. Herb. Franklin & Marshall College 1: 74, 
75. pl. 3. 

HuMsorpr, A., BoNPLAND, A. anp C. S. Kunvu. 1818. Samolus ebrac- 
teatus in Nova Gen. & Sp. 2: 223. pl. 129. 

Jounston, I. M. 1941. Gypsophily among Mexican desert plants. Jour. 
Arnold Arb. 22: 145-170. 

Mutter, C. H. 1947. Vegetation and climate of Coahuila, Mexico. Ma- 
drofio 9: 33-57. 

Pax, F. anp R. Knura. 1905. Samolus in Engler, Das Pflanzenreich 22 
(IV. 237): 336—344. 

RomiNsoN, B. L. 1892. Description of new plants collected in Mexico by 
C. G. Pringle in 1890 and 1891, with notes upon a few other species. 
Proc. Am. Acad. 27: 180. 

SMALL, J. K. 1897. Studies in the botany of the southeastern United States 
XII. Bull. Torr. Bot. Club 24: 491. 

1933. Samodia in Manual Southeastern Fl. 1028. 
1934. Samodia ebracteata in Addisonia 18: 41, 42. pl. 597. 


1958] Ehrle,—Genus Achillea in Pennsylvania 7 


A CYTOTAXONOMIC STUDY OF THE GENUS 
ACHILLEA IN PENNSYLVANIA! 


Erwoop B. EHRLE 


'The problem of speciation in the genus Achillea has received 
attention from workers at the laboratory of the Carnegie Insti- 
tution of Washington in California (Clausen, 1951; Hiesey & 
Nobs, 1952; Ehrendorfer, 1952). "These studies have been di- 
rected chiefly toward deciphering evolutionary trends and eco- 
logical specialization within this genus. Major study has been 
concentrated on forms from the Western United States and 
Europe. Very little is known of relationships among Achilleas 
of the Eastern United States. 

Herbarium specimens of eastern material, located at various 
institutions in Pennsylvania, all bear the name Achillea mille- 
folium L. The western species, Achillea lanulosa Nutt., has not 
been recognized in this state since all material of the genus 
has been somewhat automatically referred to A. millefolium L. 
Cronquist, in the new Britton and Brown Flora (Gleason, 1952), 
treats A. lanulosa Nutt. as a subspecies of A. millefolium L. and 
describes its distribution as occurring “toward the western part 
of our range” and “occasionally introduced eastward.” The 
two forms are differentiated by Cronquist on the basis of the 
degree of crowding of the leaf segments and the shape of the 
ultimate segments. 

A. millefolium L. sens. strict. is a European hexaploid with 54 
somatic chromosomes, whereas American A. lanulosa Nutt. is 
tetraploid with 36 somatic chromosomes (Ehrendorfer, 1952). 
In his review of the geographical distribution of Achillea in 
North America, Lawrence (1947) included the three eastern 
records (Vermont, Mass., & N. Y.) for tetraploid A. lanulosa 
Nutt. as reported by Turesson (1939). No cytologically proven 
records of the existence in the east of the European hexaploid, 
A. millefolium L. sens. strict., occur in the literature. 

Twenty-six specimens of Achillea were collected in Pennsyl- 
vania during the winter of 1956 and the spring of 1957 and 
were grown in a greenhouse at the Pennsylvania State University. 


l'The aid of Dr. Paul Grun in the preparation of material and in the interpretation of 
cytological figures is gratefully acknowledged. 


8 Rhodora [Vor. 60 


The leaves of these plants and those of herbarium specimens 
labeled A. millefolium or A. lanulosa from various parts of the 
United States were examined. It was not possible to separate 
the herbarium specimens or the growing plants into two categories 
on the basis of the shape of the ultimate segments or the degree 
of crowding, as the plants were highly variable with regard to 
these two characters. 

When the plants were growing vigorously, root tips were col- 
lected, treated by the oxiquinoline method (Tijo and Levan, 
1952), and stained with propio-orcein. The cells of all plants 
studied showed 36 somatie chromosomes. A list of these plants 
with their points of collection is shown in Table I. Many of the 
chromosomes (Fig. 1) were of equal length and had centrally 
located centromeres. Of those with sub-terminal centromeres, 
four satellite-bearing chromosomes were frequently discernible, 
indicating that this tetraploid species arose from satellite-bearing 
diploid precursors, either by auto- or allo-polyploidy. 

The existence of tetraploid Achillea in Pennsylvania is thus 
demonstrated. These plants should, therefore, be referred to eco- 
logical races of the western tetraploid A. lanulosa Nutt. and not 
to European A. millefolium L. as is the current practice. It might 
be possible that the latter species will be found in the vicinity 
of seaports and other points of entry of European migrants. 


TABLE I. Known distribution in Pennsylvania of tetraploid A. lanulosa 
Nutt —verified by chromosome counts. 


County Locality No. plants determined 
Centre State College 1 
Clearfield Caledonia Pike 1 
? Clearfield Bridge 3 
? Dimeling 4 
” Grampian 1 
á Kratzer Run 4 
? Lick Run 1 
Clinton Lamar 1 
Huntingdon Spruce Creek 2 
Lycoming Maple Hill 1 
Perry Mecks Corner 1 
? Millerstown 1 
” Shermandale 1 
? Waggoner Gap 1 
Snyder Kratzerville 1 
” Mt. Pleasant Mills 1 
Union Weikert 1 


1958] Ehrle,—Genus Achillea in Pennsylvania 9 


+> 
IO M 


Fig. 1. Camera lucida diagram of the mitotic early metaphase chromosomes of tetra- 
ploid Achillea lanulosa Nutt., occurring at the side of Rt. 322 at Kratzer Run in Clearfield 
Co., Pennsylvania. 


The presence of this species in such areas must, however, be 
proven before we can conclude that any A. millefolium L. sens. 
strict. exists in Pennsylvania.—DEP'T. OF BOTANY AND PLANT PA- 
THOLOGY, PENNSYLVANIA STATE UNIVERSITY, UNIVERSITY PARK, PA. 


LITERATURE CITED 


CLAUSEN, J. (1951) Stages in the Evolution of Plant Species. Cornell 
Univ. Press. Ithaca, N. Y. 

EHRENDORFER, F. (1952) Cytotaxonomic studies in Achillea. Carnegie 
Inst. Wash., Yearbook #51, pp. 125-131. 

GLEASON, H. A. (1952) The New Britton and Brown Illustrated Flora of 
the Northeastern United States and Adjacent Canada. Vol. 3, p. 385. 

Hisey, WM. M. « M. A. Noss (1952) Genetic structure of climatic 
races and species of Achillea. Carnegie Inst. Wash., Yearbook #51, pp. 
122-125. 

L^4wRENCE, WM. E. (1947) Chromosome numbers in Achillea in relation 
to geographic distribution. AJB; 34, pp. 538—545. 

Tio, J. H. ann A. Levan (1952) The use of oxiquinoline in chromosome 
analysis. Anal. de la Estacion Experimental de Aula 2, pp. 21-64. 
Turesson, G. (1939) North American types of Achillea millefolium L. 

Bot. Notiser 1939, pp. 813-816. 


10 Rhodora [Vor. 60 


SPECIES NEW TO THE FLORA OF NORTH OR 
SOUTH CAROLINA! 


Harry E. Anes, C. Rircuig BELL AND ALBERT E. RADFORD 


Most of the distribution records included in this paper are 
from the collections of the authors during 1956. Others are from 
the many excellent specimens contributed by O. M. Freeman to 
the Flora of the Carolinas project. Still others were found in the 
herbaria of the University of North Carolina (ncu), North 
Carolina State College (Ncsc), Duke University (DUKE) and the 
New York Botanical Garden (Ny). 

'The determinations of the grasses included in this paper have 
been made or verified by H. L. Blomquist of Duke University. 
R. B. Channell, then of the Gray Herbarium and Arnold Arbore- 
tum, and Joseph Monachino, of the New York Botanical Garden, 
checked their respective herbaria for the presence of many of the 
species that appear in this report. In addition, Mr. Monachino 
also verified the identification of the following specimens: Ses- 
bania exaltata (Raf.) Cory, Bell 4822; Plantago hookeriana var. 
nuda (Gray) Poe, Ahles 12169, 12607 and Bell 2712; and Hypo- 
chaeris glabra L., Ahles 12404. H. N. Moldenke supplied cur- 
rent distribution information on the verbenas. We gratefully 
acknowledge the valuable technical aid rendered by these indi- 
viduals, and the generous field assistance given us by Lionel 
Melvin, R. S. Leisner and J. H. Horton. The hospitality and 
help of the curators of the various herbaria were sincerely 
appreciated. 

All but two of the plants included in the following enumeration 
of species we presume to be new records for one or both of the 
Carolinas. The family arrangement follows that used by Fer- 
nald (1950) ; genera and species are listed alphabetically. Speci- 
mens and collection data are cited by state (North Carolina first) 
and county, the latter being arranged alphabetically. All speci- 
mens are deposited in the University of North Carolina Her- 
barium (Ncv) unless otherwise noted by an appropriate symbol. 


Andropogon campyloracheus Nash—sovrH CAROLINA. Allendale Co.: 


1JTt is a pleasure to acknowledge the generous financial assistance given in support of the 
project by Mrs. W. C. Coker; by the Alumni Annual Giving Fund and the Smith Fund 
of the University of North Carolina; and by the Highlands Biological Laboratory via a 
grant-in-aid from the National Science Foundation. 


1958] Ahles, et al.,—Species New to the Carolinas 11 
Sandhill by Co. Rt. 41 sw. of Barton and 2.1 mi. n. of jet. with Co. Rt. 
91, Bell 5124. 

Hitchcock (1933) and Chase (1950) report the range as Florida to 
Louisiana. "This specimen represents a. northern extension of range into 
South Carolina. A specimen of the cited collection was verified by 
Agnes Chase. 

Brachiaria platyphylla (Griseb.) Nash—sovrH CAROLINA. Beaufort 
Co.: Weed in vegetable garden, Dale, Ahles 15584; weed in corn field, 
0.6 mi. n. of Lobeco on U. S. 21, Ahles 20946. 

In Hiteheock (1933) this species is cited as Brachiaria extensa. Chase. 
According to Chase (1950) this grass is found primarily in the Gulf 
States with an east coast extension into Georgia. These are the first 
records of this weed from South Carolina. 

Digitaria filiformis (L.) Koel—sourH caroLINA. Hampton Co.: 
Sandy roadside, 3.1 mi. nw. of Yemassee on 8. C. 28, Ahles 15750. 

Hitchcock (1933) treats this species as Syntherisma filiforme (L.) 
Nash, giving its distribution as Florida to Texas and Massachusetts. 
Chase (1950) maps the species from all the states on the southeastern 
coast except South Carolina. No herbarium specimens have been found 
in NCU, DUKE or NCSC. 

Digitaria texana Hitche —sovuTH CAROLINA. Colleton Co.: Sandy oak 
woods at jet. of Co. Rts. 172 & 28, w. of Hendersonville, Ahles 17916. 

Hiteheock (1933) does not include this species. Chase (1950) reports 
it from southern Texas only. This is the first southeastern record for 
this taxon. 

Eragrostis lugens Nees.—sovTH CAROLINA. Hampton Co.: Pond and 
sandy margins, 0.8 mi. nnw. of Shirley on Co. Rt. 20, Bell 5013. Oconee 
Co.: Meadow near S. C. 59, 1 mi. n. of Fair Play, Radford 17916. 

Hitchcock (1933) gives the distribution of Eragrostis lugens as Lou- 
isiana to California and naturalized in Georgia and Alabama. Chase 
(1950) has essentially the same distribution. This is the first report 
of a collection from South Carolina. 

Heteropogon melanocarpus (Kll.) Benth.—sovTH CAROLINA. Beau- 
fort Co.: Sandy roadside, Hilton Head Island, Ahles 18062. Hampton 
Co.: Roadside, 1 mi. s. of Brighton, Ahles 21008. Jasper Co.: Field- 
savannah border, on S. C. 170, 1.6 mi. n. of jet. with S. C. 128, Bell 5317. 

This grass has been known previously only as far north as Georgia, 
as reported by Hiteheoek. (1933). and Chase (1950). 

Melica nitens Nutt.—NoRTH CAROLINA. Madison Co.: Hardwood 
forest, 3 mi. nw. of Hot Springs on the French Broad River, Radford 
8053. 

Chase (1950) and Fernald (1950) report the range as Pennsylvania 
to Virginia. In Madison County the species was very infrequent on 
limestone. 

Paspalum plicatulum \ichx.—souTH CAROLINA. Colleton Co.: Culti- 
vated field, Ritter, Ahles 15489. 

According to the range given for this species by Hiteheoek (1933) 


12 Rhodora | Vor. 60 


and Chase (1950), the Colleton County collection represents a northern 
extension from Georgia. 

Phalaris canariensis L.—souTH CAROLINA. Anderson Co.: Waste 
places, mixed deciduous woods, 0.5 mi. wnw. of Seneca-Tugaloo River 
jet., Radford 12087. 

This European adventive is widely scattered in waste places and 
is rarely persistent, according to Fernald (1950). Chase (1950) maps 
the species from all of the states east of the Mississippi River except 
North Carolina, South Carolina, Georgia and Alabama. The Anderson 
County collection is the first report from South Carolina. 

Scirpus etuberculatus (Steud.) Kuntze—souTH CAROLINA. Marlboro 
Co.: Fresh-water marsh, 2.25 mi. e. of Wallace on S. C. 9, Radford 
15549. 

Small (1933), Beetle (1947), Fernald (1950) and Gleason (1952) indi- 
cate the range as from Delaware to Florida and Louisiana. The absence 
of specimens from South Carolina in the NCU, DUKE, Ncsc and Ny her- 
baria warrants the inclusion of this species in the list of South Carolina 
plants. 

Scirpus koilolepis (Steud.) Gleason—NomnTH CAROLINA. Union Co.: 
Meadow, 3 mi. s. of Rocky River, 4.5 mi. n. of Unionville, Radford 
10597 ; wet corn field, 2.5 mi. w. of New Salem, Ahles 12083. 

Small (1933) treats this taxon as Scirpus carinatus (H. & A.) Gray, 
ranging from Alabama to Texas and Tennessee. Beetle (1947), Fernald 
(1950) and Gleason (1952) give essentially the same range. The 1956 
collections represent an eastward extension from Tennessee into North 
Carolina. 

Commelina communis L. var ludens (Miquel) C. B. Clarke—Norru 
CAROLINA. Cleveland Co.: Roadside ditch, 0.5 mi. n. of Lawndale on 
Maple Creek, Ahles 15195. Polk Co.: Roadside, Saluda, 6 mi. n. of 
Tryon, Fosberg 18785. 

According to Fernald (1950) this naturalized variety of the Asiatic 
species is found from New Jersey and Pennsylvania to Virginia and 
Kentucky. It is not mentioned by Small (1933). The cited specimens 
represent a southern extension of range into North Carolina. 

Allium ampeloprasum [,—NorTH CAROLINA. Davidson Co.: Road- 
side, 3 mi. nw. of Churchland near Yadkin River, Radford 12864. sovrH 
CAROLINA. Oconee Co.: Roadside of Co. Rt. 68, w. of Madison, Bell 3111. 

This Old World species is not treated by Small (1933), and Fernald 
(1950) includes it only because the var. atroviolaceum (Boiss.) Regel 
has been found in York County, Virginia (Fernald, 1941). Although 
Gleason (1952) includes the species, his treatment is apparently based 
on Fernald’s collection of the variety in southeastern Virginia. James 
(1956) reports the species in an apparently undocumented list of plants 
introduced into northern Tennessee, but without comment as to whether 
or not it is the variety mentioned by Fernald. Without herbarium 
material it is not possible to know whether James had the first. record 


1958] Ahles, et al..—Species New to the Carolinas 13 


for the species itself in the eastern United States, or another record 
for the variety. 

Brodiaea uniflora (Lindl. Engler—NomTH CAROLINA. Orange Co.: 
Field along railroad tracks at U.N.C. power plant at Chapel Hill, Rad- 
ford 8842. 

This South American horticultural introduction has become well 
established as a weed in Chapel Hill. Bailey (1950) lists the species as 
Brodiaea uniflora Baker and gives Triteleia uniflora Lindl. as a synonym; 
Hoover (1941) specifically excluded this taxon from the genus T'riteleia 
with the comment that it may belong to either Nothoscordum or 
Tristagma. 

Hymenocallis occidentalis (Le Conte) Kunth—soUTH CAROLINA. 
Union Co.: Rocky shoals of Broad River, Lockhart, Freeman 56401. 

Small (1933) states the range of this species as Georgia to Missouri 
and Indiana. Fernald (1950) gives it as Georgia to Alabama, and north 
to Kentucky, southern Indiana, southern Illinois and southeastern Mis- 
sourl. Gleason (1952) says that it occurs from southern Indiana to 
southern Missouri and south to the Gulf. Apparently the Union County 
collection is the first record for South Carolina. 

Pilea fontana (Lunell) Rydb—NorrH CAROLINA. Halifax Co.: 
Swamp along the Roanoke River, just n. of Roanoke Rapids on N. C. 
48, Ahles 20713. Jones Co.: Hardwood forest over marl on Trent River, 
ne. of Pollocksville, Radford, Haesloop & Miller 7707. 

The two preceding specimens are recorded here because of the dis- 
crepancy in the ranges given by Fernald (1950), “w. N. Y. to N. D., 
s. to n. Fla., Ind., Ill. and Neb.,” and Gleason (1952), “P. E. I. to Minn. 
& N. D. s. to Va., Ind., and Neb.; Fla.” Small (1933) does not include 
this species at all. The placing of Pilea opaca (Lunell) Rydb. as a 
synonym of this species in Gleason (1952) rather than as a synonym 
of P. pumila (L.) Gray is apparently an error, as is the illustration for 
this species which is clearly Boehmeria cylindrica (L.) Sw. However, 
the achene to the left of the illustration is that of P. fontana. 

Polygonum hirsutum Walt.—sovTrH CAROLINA. Allendale Co.: Low 
pine savannah, 1.5 mi. se. of jet. Co. Rts. 41 & 60 on Co. Rt. 26, sw. 
of Barton, Ahles 15868; pond banks, 7 mi. nw of. Allendale on S. C. 
641, Bell 4039; field, 3.3 mi. sw. of Sycamore, 3 mi. e. of jet. U. S. 301 
& S. C. 644, Bell 5185. Lexington Co.: Mucky margin of open shallow 
pond, Columbia, Godfrey & Tryon 1220 (DUKE, NY). 

All of the plants described by Walter in his Flora Caroliniana (1788) 
are presumably from South Carolina. Apparently Small (1895) over- 
looked this fact or had reason to ignore it, in giving the range of Poly- 
gonum hirsutum as Georgia and Florida. In his 1933 manual he repeats 
the same range under the name Persicaria hirsuta (Walt.) Small. Due 
to the uncertainty of the existenee of an herbarium specimen from 
South Carolina, we are citing the above collections. 

Rumex patientia L.—NorTH CAROLINA. Wake Co.: Waste ground, 
Rolesville, Ahles 11428. 


14 Rhodora [Vor. 60 


Small (1933) does not include this species. Fernald (1950) reports 
it south to eastern Pennsylvania while Gleason (1952) gives no indi- 
cation that it occurs outside of that manual’s range. Therefore the 
preceding specimen is cited as the first record for North Carolina. Only 
two plants were observed at the Rolesville locality. 

Cycloloma atriplicifolium (Spreng.) Coult.—NonTH CAROLINA. New 
Hanover Co.: Sandy waste place, N. C. 420 at causeway, Wrightsville 
Beach, Radford & Radford 2079. SOUTH CAROLINA. Darlington Co.: 
Common in a field on road from Darlington to Lauther's Lake, B. E. 
Smith 1657 ; sandy soil on Witherspoon Island, B. E. Smith 916. Sum- 
ter Co.: Roadside ditch 1 mi. n. of county line on U. S. 15A, Bel! 3607. 

These collections are recorded here because the Southeastern United 
States are excluded from the range of this taxon as given by Small 
(1933), Fernald (1950) and Gleason (1952). At the Sumter County 
locality the species was abundant in several fields. 

Amaranthus graecizans L.—sovTH CAROLINA. Anderson Co.: Field, 
S. C. 81, 3.5 mi. n. of Anderson, Radford 14053. 

Although the ranges given by Small (1933), Fernald (1950) and 
Gleason (1952) indicate that this species occurs throughout the United 
States, intensive field work in both North and South Carolina during 
1956 produced only one collection. No specimens from either state are 
present in the herbaria of NCU, DUKE, Ncsc or Ny. These facts seem 
to warrant the reporting of the Anderson County collection as a new 
record for South Carolina. 

Froelichia gracilis (Hook.) Moq.—sovrH CAROLINA. Allendale Co.: 
Sandy field sw. of Allendale on Co. Rt. 57, Bell 4034. Lee Co.: Sandy 
waste ground, 1 mi. sw. of Aleot on U. S. 15, Ahles 15898. 

According to Fernald (1950) and Gleason (1952), this species is native 
to the central United States and is adventive eastward, ranging south- 
ward only as far as Virginia. Small (1933) makes no mention of it. 
Froelichia gracilis is probably more widely distributed throughout the 
Southeastern United States than present collections would indicate, as 
it is easily confused with F. floridana (Nutt.) Moq. and possibly has 
been mistaken for that species. However, the gray color, low stature 
and habit of branching freely at the base distinguish F. gracilis from 
F. floridana. Froelichia gracilis is to be expected along railroads or 
in sandy places. 

Mirabilis jalapa L.—NonrH CAROLINA. Iredell Co.: Statesville M. £F. 
Hyams 4991. Stanly Co.: Well established on road embankment, 4.1 
mi. sw. of N. C. 73 on N. C. 27, Ahles 19859. 

Small (1933) says of this species, “escaped, s. Fla.,” while Fernald 
(1950) and Gleason (1952) give a very vague range. In 1956 it was 
found well established on a road embankment and for this reason 1s 
placed on record. 

Tetragonia expansa Murr.—NonTH CAROLINA. Durham Co.: Escaped 
from gardens and fields, Engelwood Ave., Durham, Blomquist 3584 
(DUKE). 


1958] Ahles, et al.,—Species New to the Carolinas 15 


Small (1933) gives the range as Florida and California. Fernald 
(1950) says, “sometimes escapes from cultivation,” and Gleason (1952) 
states, “occasionally persists in gardens or appears spontaneously near 
them.” In North Carolina this species is probably no more than a 
waif, occurring sporadically after cultivation. 

Holosteum umbellatum L.—souTH CAROLINA. Cherokee Co.: Road- 
side, 8.7 mi. nw. of Gaffney on S. C. 11, Ahles 11291. Spartanburg Co.: 
Anderson Bridge over the Enoree River, S. C. 417, Freeman 6509. 

Maguire (1952) gives the range as eastern Massachusetts to Georgia 
in the Atlantic States. Even though South Carolina is in the general 
range of this species, no South Carolina specimens have been found in 
NCU, DUKE, NY Or GH. 

Spergula pentandra L.—NoRTH CAROLINA. Warren Co.: Sandy road- 
side, 0.5 mi. e. of Warren Plains on U. S. 158, 200 yards e. of railroad 
crossing, s. side of highway, Ahles 10966. 

According to Maguire (1952) this European species is an adventive 
at Cape May, New Jersey only. The Warren County collection is the 
first record for the southeast. 

Ranunculus arvensis L. var. tuberculatus DDC.—NORTH CAROLINA. 
Cabarrus Co.: Fields, Harrisburg, May 25, 1944, W. H. Garmon (Ncsc) ; 
field by woodland, 3 mi. ne. of Harrisburg, along Wolf Meadow Branch, 
Bell 2271. Clay Co.: Roadside, 1.3 mi. sw. of Hayesville at jet. of 
Sweet Water Gap Road and U. S. 64, Ahles 13801. Mecklenburg Co.: 
Wheat field, 1.5 mi. w. of Mecklenburg-Cabarrus Co. line on N. C. 49, 
Ahles 11324. 

Gleason (1952) describes the species as a European introduction 
occasionally found along the Atlantie coast. The variety is not described 
by Small (1933), Fernald (1950) or Gleason (1952). According to Hegi 
(1912) it is found in Central Europe. In North Carolina the variety 
seems to have been introduced with grain, and therefore may be estab- 
lished elsewhere in North Ameriea. At one of the localities the species 
grew mixed with the variety, and at two localities the variety occurred 
by itself. This variety of R. arvensis was very abundant at two of 
the localities. 

Ranunculus sardous Crantz—NorTH CAROLINA. Iredell Co.: Road- 
side, 2.9 mi. w. of Elmwood and 1.5 mi. ne. of U. S. 70 near Fourth 
Creek, Ahles 12982. Union Co.: Roadside, 0.3 mi. s. of Stanly-Union 
Co. line (Rocky River) about 4.5 mi. n. of Unionville, Ahles 11952. 
SOUTH CAROLINA. Colleton Co.: Wet soil near railroad, Ritter, Ahles 
12177. 

Small (1933) gives the range of this species, under Ranunculus 
parvulus L., as *Cosatal Plain and New England Coast, Ga. to N. B." ; 
Fernald (1950) says, “N. B. to Mo. and southw." According to Gleason 
(1952) this weed ranges from southeastern Virginia to Georgia. No 
specimens from either of the Carolinas have been found in the local 
herbaria. 

Alliaria officinalis Andrs.—NORTH CAROLINA. Rockingham Co.: 


16 Rhodora [Vor. 60 


Wooded river bottom, about 2 mi. e. of Thompsonville on N. C. 87, 
Ahles 10746; swamp forest, Dan River near N. C. 14, s. of Leaksville, 
Radford 10769. 

Small (1933) does not include this species in his manual. Fernald 
(1950) and Gleason (1952) have it ranging only as far south as Virginia 
on the eastern portion of the range. 

Hesperis matronalis L.—NonTH CAROLINA. Forsyth Co.: Escape, 
Winston-Salem, Schallert 8765. 

Although the range of this species as given by Small (1933), Fernald 
(1950) and Gleason (1952) may be interpreted to include North Caro- 
lina, there are no specimens from the state at DUKE, Ncsc or NY. This 
fact warrants the inclusion here of the Forsyth County record. 

Raphanus raphanistrum L.—souTH CAROLINA. Beaufort Co.: Road- 
side and edge of cultivated field, Bluffton, Ahles 12372. Darlington Co.: 
Field s. of ferry at Lauther’s Lake, B. E. Smith 1492. Hampton Co.: 
Sandy roadside, 2 mi. n. of Estill on U. S. 321, Ahles 10588; weed in 
cultivated field, 3.7 mi. nne. of Early Branch, Ahles 12/62. Jasper Co.: 
Waste ground, Ridgeland, AAles 15607. Marlboro Co.: Shrub bog, 2.5 
mi. nw. of Blenheim, Radford 12667. 

This species is not included in Small (1933), and is recorded only as 
far south as Virginia in the eastern part of the ranges as given by Fernald 
(1950) and Gleason (1952). Freeman (1955) reports it from North 
Carolina and Thorne (1951) from Georgia. The Jasper County speci- 
men, Ahles 15607, is forma albus (Schuebler & Martens) Hayek. 

Thlaspi perfoliatum L.—NonrH CAROLINA. Alexander Co.: Field, 
4 mi. e. of Caldwell Co. line on N. C. 90, Radford 9979. Graham Co.: 
Roadside, about 2 mi. s. of the town of Yellow Creek on U. S. 129, Ahles 
18317. Rockingham Co.: Meadow, 3 mi. n. of Reidsville near N. C. 14, 
Radford 13612. Stokes Co.: Roadside, 0.4 mi. w. of Stokes-Rockingham 
Co. line on U.S. 311, about 1 mi. n. of Pine Hall, Ahles 10807. 

Thlaspi perfoliatum is not included by Small (1933), and is reported 
south only to Virginia by Fernald (1950) and Gleason (1952). 

Warea cuneifolia (Muhl.) Nutt.—NoRTH caRoLiNA. Harnett Co.: 
From sandhills, about 3 mi. ne. of Spout Springs on Sprunt land, 
Sept. 1, 1940, H. A. Rankin. 

Small (1933) gives the range as “Fla. to Ala. and S. C.” The Harnett 
County collection is not a new record as it was cited in 1940 by Rankin 
in the Journal of the Gray Memorial Botanieal Association. However, 
due to the omission of a complete citation of Rankin’s specimen in 1940 
in an obscure publication, it would seem worthwhile to repeat this 
state record here. 

Chrysobalanus oblongifolius Michx.—sovTH CAROLINA. Jasper Co.: 
Sand hill, 2 mi. nw. of Grays on S. C. 631, Ahles 15794; sand hills, 10.5 
mi. nw. of Tillman, 0.6 mi. se. of Seaboard Air Line Railway, AAles 18217. 

Small (1933) gives the range of this species, under the name Geo- 
balanus oblongifolius (Michx.) Small, as “Fla. to Miss. and Ga.” The 


1958] Ahles, et al.,—Species New to the Carolinas 17 


species was found to be fairly abundant in the sand hill country of 
northwestern Jasper County where it appears to be native. 

Sesbania exaltata (Raf.) Cory— NORTH CAROLINA. Brunswick Co.: 
Abundant in extensive marsh, Brunswick River w. of Wilmington, 
Godfrey 50162 (DUKE, Ncsc). Granville Co.: Camp Butner, Batson 
118 (DUKE). SOUTH CAROLINA. Darlington Co.: Coker's Seed Farm, 
Hartsville, B. E. Smith 1559. Jasper Co.: Sandy roadside, edge of 
brackish marsh, 0.7 mi. n. of the Savannah River on U. S. 17A, Ahles 
18154. 

The range of this species as given by Small (1933) under Sesban 
exaltata (Raf.) Rydb. does not include the South Atlantic States. 
Thorne (1951) reports it for the first time from Georgia while the above 
earlier and unpublished collections from North and South Carolina 
were found during the checking of herbarium specimens in connection 
with the current collection from Jasper County. 

Oxalis martiana Zucc.—NORTH CAROLINA. Wake Co.: Roadside near 
Cary, April 1, 1949, Godfrey (Ncsc). 

Small (1933) under Jonozalis martiana (Zuce.) Small, gives the range 
of this species as “Fla. to Tex. and S. C." No specimen from North 
Carolina has been found in Ncv, DUKE or Ny. Therefore, the above 
colleetion is cited here as the first record for North Carolina. 

Ruta graveolens L.—NonTH CAROLINA. Orange Co.: Roadside on 
edge of field by Morgan Creek, above bridge on Pittsboro Rd., Chapel 
Hill, April 8, 1945, Costello & D. & A. Beers. 

Small (1933) gives the range for this species as various provinces in 
the eastern United States. Gleason (1952) cites no range, merely stating, 
"Native of Europe; cultivated in old-fashioned gardens and often 
escaped to waste ground and roadsides.” Fernald (1950) gives it south 
only to Virginia on the Atlantic Seaboard. The Orange County col- 
lection is cited here as a first record for North Carolina. 

Croton lindheimeri (Engelm. & Gray) Wood—souTH CAROLINA. 
Beaufort Co.: Sandy palm woods, Jenkins Island, Bell 4807. Richland 
Co.: Vacant lot near railroad, Columbia, Freeman 56823. 

According to the range given by Small (1933) for this species under 
the binomial Croton engelmannii Ferguson, and that given by Fernald 
(1950), the Beaufort and Richland County collections represent a 
northern extension in the east coast states; both Small and Fernald 
report it north only to Georgia on the Atlantic Seaboard. Gleason 
(1952) treats this entity as C. capitatus var. lindheimeri Muell.-Arg., 
apparently ignoring the authority (Engelm. & Gray) for the basionym, 
and reports it only in Florida in the Atlantic coast states. At the Beau- 
fort County locality it was very abundant, and conspicuous from a car. 

Euphorbia dentata Michx.—NoRTH CAROLINA. Graham Co.: Road- 
side, U. S. 129, near Cheoah Creek, 5 mi. se. of Tapoco, Radford 15904. 
Surry Co.: Ditch, Yadkin River near Elkin, Radford 18380. 

Small (1933), under Poinsettia dentata (Michx.) Small, has this 
species as occurring throughout the southeastern United States, while 


18 Rhodora [Vor. 60 


Gleason (1952) excludes this region, at least south of Virginia. In 
Fernald (1950) the range “N. Y. to Minn., S. D. and Wyo., s. to Va., 
La., Tex. and Mex.,” can be interpreted as either including or excluding 
North Carolina. The absence of specimens in NCU, DUKE and NCSC is, 
therefore, the principal reason for including this species as new to 
North Carolina. 

Euphorbia hirta L.—souTH CAROLINA. Jasper Co.: Lawn weed 
around gas station, Ridgeland, Ahles 15812. 

Small (1933) under the name Chamaesyce hirta (L.) Millsp. gives 
the range of this species as “Fla. to Tex." Fernald (1950) does not 
mention it at all and Gleason (1952) says, “Widely distributed in tropical 
and subtropical lands, including the southern states.” Since no mention 
is made of South Carolina, the specimen from Jasper County is worthy 
of note here as the species is well established at this locality. 

Phyllanthus niruri L.—sovTH CAROLINA. Sumter Co.: Weed in dis- 
turbed soil, Sumter, Freeman 56801. 

'The range given by Small (1933) for this southern species is Florida 
and the Keys. It spreads freely by seed as a weed in the greenhouse 
of the University of North Carolina. For this reason it is not too sur- 
prising to find it spreading, particularly in disturbed areas. More careful 
field work will doubtless reveal it in other places. 

Phyllanthus pentaphyllus C. Wright—sourH CAROLINA. Darlington 
Co.: In sand hills bevond Camden Rd., 5 mi. n. of Hartsville, March 
27, 1910 (without collector or number). 

This species is known only from Florida, the lower Florida Keys and 
the West Indies, according to Small (1933). It is possibly only a 
sporadie introduction in South Carolina, or it may be an isolated 
native remnant. 

Stillingia aquatica Chapm.—souTH CAROLINA. Hampton Co.: Asso- 
ciated with Taxodium ascendens Brongn. and /lexr myrtifolia Walt., 0.2 
mi. n. of Luray on U. S. 321, Ahles 18285. 

Small (1933) gives the range of this species from Florida to Mississippi 
and South Carolina. However, Rogers (1951) restriets its distribution 
to Florida, and Sumter County, Georgia. The fact that UNC, DUKE, 
Ncsc and NY have no specimens other than those from Florida and 
Georgia is significant. The Hampton County specimen is considered 
a northward extension of the range. 

Callitriche terrestris Raf—NorrH CAROLINA. Union Co.: Meadow, 
3 mi. s. of Rocky River, 4.5 mi. n. of Unionville, Radford 10598. Warren 
Co.: Wet border of cultivated field, about 4 mi. n. of Church X Roads 
on Jordans Creek, near the Roanoke River, AAles 12865. 

Small (1933) and Fassett (1951) give the range of this species as 
western Massachusetts south to Virginia, and in the midwest south to 
Louisiana and Alabama. The Union and Warren County collections 
are a southern extension of range into North Carolina. 

Elaeagnus pungens Thunb.—NoRTH CAROLINA. Forsyth Co.: Escape, 
Schallert 613 (no locality). Orange Co.: Pine-oak woods, Chapel Hill, 


1958] Ahles, et al.,—Species New to the Carolinas 19 


March 10, 1956, Ahles 9995. souTH CAROLINA. Union Co.: S. C. 9 
at the Broad River, Freeman 56880. 

This commonly cultivated species is not included by Small (1933), 
Fernald (1950) or Gleason (1952). James (1956) seems to be the first 
to have reported it as established in a southeastern state. It occurs 
abundantly in small colonies in each of the localities listed. It is quite 
frequent in the woods around Chapel Hill as an escape. 

Cuphea carthagenensis (Jacq.) Macbr.—NonTH CAROLINA. Brunswick 
Co.: Swampy woods along creek, Orton, Godfrey 10156. Carteret Co.: 
Wet sandy soil, Beaufort, July 15, 1938, R. Gray (DUKE, Ncsc); pine- 
land at Sea Level, Godfrey 6412. Johnston Co.: Savannah along U. S. 
70 near Pine Level, Radford 4409. Onslow Co.: Roadside at Folk- 
stone, July 25, 1923, E. J. Alexander. Pitt Co.: Path near house site, 
1 mi. e. of Fountain, Radford, Haesloop & Miller 7357. Wayne Co.: 
Marshy ground at edge of lake, about 5 mi. ne. of Mt. Olive, Blomquist 
14768 (DUKE). Wilson Co.: Pineland, 12 mi. e. of Wilson, Blomquist 
11214 (DUKE). SOUTH CAROLINA. Beaufort Co.: Swamp 0.2 mi. ne. of 
Neue River on S. C. 170, Bell 5283. Colleton Co.: Roadside swamp, 
0.7 mi. sw. of Cottageville, on Co. Rt. 40, AAles 17949; under highway 
bridge over Combahee River on U. S. 17A, Ahles 21031. Georgetown 
Co.: Sand wash along small stream 2 mi. ne. of Andrews, Godfrey & 
Tryon 964 (DUKE). Jasper Co.: Swamp by 8. C. 170, 2 mi. s. of jet. 
with S. C. 128, Bell 4837. 

Small (1933) gives the range of this plant, under the name of Par- 
sonsia balsamona (C. & S.) Standley, as “lake region, pen. Fla." Thorne 
(1951) reported the species from Georgia. It was found in both North 
and South Carolina during the 1956 collecting season. The earlier 
specimens cited above were found while checking the local herbaria; 
many were misidentified, or identified only to genus. Despite the 
relatively large number of these collections from the Coastal Plain, this 
species does not appear to have been published previously for North 
or South Carolina. Macbride (1930) pointed out that Lythrum 
carthagenense Jacq. and Cuphea balsamona Cham. & Schlecht. were 
names applied to the same species; therefore, he made the new combi- 
nation now in use. 

Gaura parviflora Dougl—souTH CAROLINA. Aiken Co.: Near Sa- 
vannah River on U. S. 1, Radford 513. Fairfield Co.: Railroad crossing 
at Dawkins, Freeman 56395. 

Small (1933) reports the range of this taxon as Alabama and west- 
ward. Munz (1938) gives the range of the species and the varieties as 
mid-western and western. The Aiken and Fairfield County collections 
are the first records for South Carolina. 

Jussiaea leptocarpa Nutt.—NoRTH CAROLINA. Harnett Co.: Pond 
margin, 10.6 mi. e. of Harnett-Lee Co. line on U. 5S. 421, H. Laing 625. 
SOUTH CAROLINA. Beaufort Co.: Brackish marsh, 0.7 mi. sw. of Co. 
Rt. 33 on U. S. 17, Ahles 18015. Hampton Co.. Sandy shore of pond, 
0.8 mi. nnw. of Shirley on Co. Rt. 20, Ahles 18257. Jasper Co.: Edge 


20 Rhodora [Vor. 60 


of eypress swamp, 2.4 mi. s. of jet. of U. S. 17 & 17A on U. S. 17A, 
Ahles 18149. Lee Co.: Near U. S. 15, about 1 mi n. of DuBose, Free- 
man 6788. 

Small (1933), Fernald (1950) and Gleason (1952) give the range of 
this plant as Florida to Texas, and north to Georgia in the Atlantic 
States. The cited specimens extend the range northward to South and 
North Carolina. 

Anthriscus scandicinus (Weber) Mansf.—NoRTH CAROLINA. Madison 
Co.: Waste ground at Hot Springs, Freeman 56330. 

This European weed is not mentioned by Small (1933) or Rodgers 
(1950). Fernald (1950) states that it is “as yet local" in southeast 
Virginia. 

Chaerophyllum procumbens (L.) Crantz—NorTH CAROLINA. Ala- 
mance Co.: Wooded floodplain of Haw River, 11 mi. w. of Chapel Hill, 
Radford 5572. Lee Co.: Woods on floodplain of Cape Fear River near 
Hgih Hill, Radford 5559. Rockingham Co.: River bottom woods along 
Dan River, 1.5 mi. ese. of Leaksville, Ahles 10780. Warren Co.: Wooded 
slopes of river bottom by Jordan's Creek at Roanoke River, about 4 
mi. n. of Church X Road, Bell 2860. 

Although this species was listed by Curtis (1867) as growing in the 
"Mid. Dist." (Piedmont) of North Carolina, the manuals currently 
covering our area, as do Mathias and Constance (1944-45), give the 
range of this species as generally north and west of the Carolinas. 
Rodgers (1950) cites two nineteenth century collections from South 
Carolina, but speaking of North Carolina he says, “No collection is yet 
known from this state." Because the Curtis reference to Chaerophyllum 
procumbens has been either overlooked or, for lack of herbarium speci- 
mens, discounted by all contemporary workers, the species is here con- 
sidered as new to the state, and the above collections are cited to show 
the plant to be growing at several points in the Piedmont of North 
Carolina. The determination of the Alamance County collection was 
verified by Dr. Lincoln Constance. 

Hydrocotyle sibthorpioides Lam.—souTH CAROLINA. Darlington Co.: 
Lower lawn of Kalmia Gardens, Hartsville, Bell 1574. 

Mathias and Constance (1944-45), Fernald (1950) and Gleason 
(1952) variously report the sporadic introduction of this small plant 
into the general area of Pennsylvania, Virginia, Kentucky and Indiana. 
This species is not mentioned by Small (1933), but specimens from 
North Carolina, Georgia and Louisiana are cited by Rodgers (1950), 
and an additional North Carolina station for the species is reported by 
Freeman (1955). No previous report of this plant from South Carolina 
is known. 

Ptilimnium fluviatile (Rose) Mathias—NonTH carRoLINA. Chatham 
Co.: In gravelly, seeping soil; rocky bed of Deep River 1 mi. w. of 
Moneure below Lockville Dam, Beard 1277. Granville Co.: Wet sand, 
growing in sand bar in middle of Tar River 10 mi. s. of Oxford on 
N. C. 96, Gillespie 397 (Ncsc). 


1958] Ahles, et al..—Species New to the Carolinas 21 


Small (1933) and Mathias and Constance (1944-45) treat this species 
as an Alabama endemic; Small uses the name Harperella fluviatilis Rose. 
Rodgers (1950) does not include the species for North Carolina. The 
first collection for this state was made by Beard while he was working 
on the flora of the Deep River basin. 

Buddleia lindleyana Fort.—NonTH CAROLINA. Forsyth Co.: Escaped 
near Winston-Salem, Schallert 508. 

Small (1933) under the name Adenoplea lindleyana (Fort.) Small 
gives “Fla. to Tex. and Ga.” as the range for this species. In 1948 E. J. 
Alexander, H. E. Ahles and J. K. McGrath collected it as established in 
the vicinity of Jocassee, Oconee County, South Carolina. A specimen 
of this collection is in the personal herbarium of H. E. Ahles. Schallert’s 
specimen appears to be the first collection for North Carolina. It is not 
known with any certainty that the plant is established in this state. 

Centaurium umbellatum Gilib.—NorTH CAROLINA. Orange Co.: Lawn 
of President's Mansion, Chapel Hill, July 6, 1914, W. C. Coker. 

Small (1933) does not include this species in his manual. Fernald 
(1950) says, “local, Que. to Mich., s. to Ga. and Ind." and Gleason 
(1952), “N.S. to n. Ill.” Because the limits of the range of this species 
are indefinite and because the Coker specimen is the only one from 
North Carolina found in any of the four herbaria, NCU, DUKE, NCSC 
or NY, the Orange County collection is considered to be the first record 
for North Carolina. It is probably only a sporadic waif in this state. 

Ampelamus albidus (Nutt.) Britt.—NonTH CAROLINA. Halifax Co.: 
Climbing on other plants, roadside, Weldon, AAles 14831; climbing in 
thickets along the Roanoke River just n. of Roanoke Rapids on N. C. 48, 
Ahles 20732. Warren Co.: Fence row and roadside at Eaton Ferry, 
Roanoke River, s. of Elams, Bell 4363. 

Although Small (1933) under Gonolobus laevis Michx. states the 
range of this plant as “Fla. to Tex., Kans., and Pa.,” Fernald (1950) 
and Gleason (1952) give the range in such a way as to exclude Florida 
and the Carolinas. 

Phlox drummondii Hook.—sovrH CAROLINA. Allendale Co.: Road- 
side about 2 mi. ese. of Ulmers on Co. Rt. 48, Ahles 12578. Beaufort Co.: 
Sand and shell dunes by U. 8. 21, near Harbor River, Bell 1723. Colleton 
Co.: Roadside and low woods, Buckhead Creek along Co. Rt. 48, 3.4 mi. 
sw. of U. S. 21, Bell 2286. Hampton Co.: Roadside of U. S. 601 about 2 
mi. nw. of Miley, Bell 2564. Jasper Co.: Roadside 2 mi. s. of Tillman 
on U. S. 321, Bell 1663. 

This plant, native to the southwest, is considered by Small (1933) to 
be naturalized east to Florida and Georgia. It is now well established 
in some parts of South Carolina. 

Heliotropium amplexicaule Vahl—NonTH CAROLINA. Anson Co.: 
Sandy pine-oak woods border, 1.6 mi. ese. of Cason Old Field, Ahles 
19471. 

Small (1933), under the name Cochranea anchusaefolia (Poir.) 
Guerke, gives the northernmost range of this plant as Georgia. It has 


22 Rhodora [Vor. 60 


been reported since for South Carolina by Smith (1946). Fernald (1950) 
gives the range of this species as “Fla. to Tex. and Calif. n. to N. J. and 
O." Gleason (1952) says, "S. C. to Fla. and Tex. occasionally n. to Va. 
and Mo." The lack of specimens from North Carolina in the local her- 
baria warrants the reporting of the Anson County collection. 

Myosotis versicolor (Poir.) Sm.—NORTH CAROLINA. Granville Co.: 
Field at Creedmoor, Radford 10449. 

Small (1033) does not mention this species, and Fernald (1950) cites 
it only as far south as Virginia where he collected it in Dinwiddie County 
(Fernald, 1938). It appears that there are no collections from North 
Carolina prior to 1956. 

Verbena brasiliensis Vell —soutH CAROLINA. Beaufort Co.: Railroad 
in Yemassee just s. of Co. line on U. S. 17A, Bell 2520. Cherokee Co.: 
Roadsides and fields, 3 mi. ne. of Chesnee, Freeman 56425. Chester Co.: 
Roadside, 0.25 mi. e. of Lockhart, Freeman 56406. Hampton Co.: Road- 
side, Yemassee, Bell 2595. Marlboro Co.: Roadside, Crooked Creek, 5 
mi. sw. of Bennettsville, Radford 15482. Sumter Co.: U. 8. 76 at the 
Wateree River, Freeman 56809. Union Co.: Waste places, Lockhart, 
Freeman 56402. 

Small (1933) gives the range of this species as Louisiana, Since that 
time it has apparently become more widely spread. Fernald (1950) says, 
“Fla. to La. and n. to se. Va.,” while Moldenke (1952) has a similar 
range, but extending it westward to Arkansas and also into southern 
California. Moldenke (1949) does not include this species for South 
Carolina. 

Verbena X engelmannii \[oldenke—NorTH CAROLINA. Ashe Co.: 
Banks of New River, Solitude P. O. (without date, collector or number). 

This hybrid of Verbena hastata L. X Verbena urticifolia L. is not 
included by Small (1933) or Fernald (1950). Moldenke (1952) cites 
this hybrid from fourteen states; none, however, are southeastern except 
Tennessee and Kentucky. In his 1949 treatment he cites no records for 
North Carolina. 

Verbena X hybrida Voss—sovrH CAROLINA. Darlington Co.: Low, 
sandy bottom lands, oper woods, se. of Hartsville, April 8, 1921, J. B. 
Norton. 

This common garden hybrid is not listed in Small (1933) or Fernald 
(1950). Moldenke (1952) says, “naturalized, Pa., O., Ia., Okla., occa- 
sionally found escaped elsewhere,” and in his treatise of this family 
(1949) he does not cite any specimens of the hybrid from South Carolina. 

Verbena rigida Spreng.—sovTH CAROLINA. Allendale Co.: Sandy 
roadside, 2.6 mi. nnw. of Allendale on S. C. 28 and 3.8 mi. ene. on Co. 
Rt. 39, Ahles 12581. Beaufort Co.: Forming colonies on roadside, 1.6 
mi. se. of Pocotaligo on U.S. 21, Ahles 12368. Colleton Co.: Road em- 
bankment, 1.4 mi. ne. of Ashepoo on U. S. 17, Ahles 15505. Dorchester 
Co.: Open sandy soil in and around Summerville, Correll 5348 (DUKE). 

Small (1933) gives the range of this introduced species as “W. Fla. to 
Tex. and N. C.” Moldenke (1949) cites many specimens from Georgia, 


1958] Ahles, et al.,—Species New to the Carolinas 23 


only a few from North Carolina, but none from South Carolina. Al- 
though South Carolina is well within the range of the species, the fact 
that Moldenke has not seen any specimens from that state warrants the 
inclusion of the collections cited above. 

Perizoma rhomboidea (Hook.) Small—NorrH CAROLINA. Carteret 
Co.: By Beaufort Channel, Beaufort, Rogers & Blomquist 3168 (DUKE). 
Orange Co.: Weed in arboretum of U. N. C., Chapel Hill, Ahles 16310. 
Rowan Co.: Abundant weed in cultivated field near Salisbury, Oct. 12, 
1955 (sent in by County Agent), (Ncsc) ; abundant garden weed, Spen- 
cer, Ahles 14933. SOUTH CAROLINA. Beaufort Co.: Along sidewalk, 
Hamar St., one block s. of Green St. (two blocks s. of National Ceme- 
tery), Beaufort, Ahles 20926. Newberry Co.: Near Wallace Home on 
Caldwell St., Newberry, Freeman 56633. 

Small (1933) gives only Florida as the range of this plant which has 
been introduced into the southeast. It now appears to be well established 
at the North Carolina localities listed above, and also in the Piedmont 
and Coastal Plain of South Carolina. The two collections from Rowan 
County are from the same locality, where the plant has become an un- 
desirable weed difficult to eradicate. 

Ajuga reptans L.—NORTH CAROLINA. Orange Co.: Growing in bed 
of Siberian iris near bamboo colony in arboretum, Chapel Hill, L. M. 
Radford & E. Wicker; March 28, 1945, W. Webb; April 17, 1947, B. 
Ivey 82; May 3, 1949, P. Titman. Wake Co.: Campus, N. C. State 
College, Raleigh, April 13, 1943, Wells (wesc); April 2, 1948, Williamson 
(Ncsc). 

Small (1933) makes no mention of this species. Fernald (1950) says, 
"Nfld., to Wisc. s. to Pa. & O.,” and Gleason (1952) states, “rarely es- 
caped near gardens." In Chapel Hill it has become fairly well estab- 
lished. The Orange and Wake County collections are being cited as the 
first report of this species for North Carolina. 

Dicerandra densiflora Benth.—sovTH CAROLINA. Jasper Co.: Sandy 
hills, lightly wooded, 7.9 mi. nw. of Tillman, 3.2 mi. se. of Seaboard Air 
Line Railway, AAles 18205; also Oct. 11, 1956, AAles 20988. 

The range of this southern species is given by Small (1933) as “e. 
Fla. and Ga." In northwestern Jasper County this plant is very local 
on the same sand ridge as Chrysobalanus oblongifolius, and is probably 
also native here. The specimen was just coming into flower on Septem- 
ber 9, and was in fruit on October 11. 

Stachys sieboldii Miq.—NoRTH CAROLINA. Cumberland Co.: Escaped 
weed in Butler’s Nursery, July 2, 1951, Butler & Totten. SOUTH CARO- 
LINA. Oconee Co.: Roadbank and upland woods by Co. Rt. 68, 2.4 mi. 
se. of Co. Rt. 17, w. of Madison, Bell 3109. 

Small (1933) and Gleason (1952) do not include this species in their 
manuals, and Fernald (1950) does not give a range, merely stating that 
it spreads from cultivation but is hardly established. At the Oconee 
County locality this species is well established on a road bank. 


24 Rhodora [ Vor. 60 


Verbascum virgatum Stokes—NorrH CAROLINA. Moore County: 
Sandy roadside, 1 mi. s. of Aberdeen on U. S. 15-501, Ahles 15377. 

Small (1953) gives South Carolina as the range for this speeies in the 
Atlantic States. Although Fernald (1950) says the range extends from 
Ontario to South Carolina, no specimens from North Carolina have been 
found in the local herbaria or in the Gray Herbarium prior to the Moore 
County collection. 

Scoparia dulcis L.—sourH CAROLINA. Colleton Co.: Under highway 
bridge, along Combahee River on U. S. 17A, Ahles 21030. 

This species has been reported by Pennell (1933) as far north as 
southern Georgia. It has not been reported previously for South Car- 
olina. 

Plantago hookeriana F. & M. var. nuda (Gray) Poe—NorTH CAR- 
OLINA. Moore Co.: Along railroad at U. S. 1 in Aberdeen, Bell 2712. 
Richmond Co.: Roadside 3 mi. ne. of Hamlet on N. C. 77, Bell 2711. 
SOUTH CAROLINA. Allendale Co.: Roadside of Co. Rt. 21 about 2.5 mi. 
sw. of Fairfax, Bell 2648. Bamberg Co.: Abundant on roadside with 
Plantago aristata Michx., 0.9 mi. ne. of Bamberg-Allendale Co. line 
(Salkehatehie River) on U.S. 301, Ahles 12605. Calhoun Co.: Roadside, 
1.6 mi. s. of Calhoun-Richland Co. line (Congaree River) on U. S. 601, 
Ahles 12608. Chesterfield Co.: Roadside, 3.6 mi. ene. of Chesterfield- 
Kershaw Co. line (Lynches River) on U. S. 1, 1 mi. sw. of McBee, Ahles 
12612. Colleton Co.: Cinder bed along railroad, Lodge, Ahles 12146; 
cinder bed along railroad, Green Pond, on 8. C. 303, Ahles 12169. Hamp- 
ton Co.: Associated with Plantago aristata Michx., roadside, 1.8 mi. nw. 
of Yemassee on S. C. 28, Ahles 12412. Kershaw Co.: Roadside, 0.5 mi. 
s. of jet. U. S. 601 and 8. C. 12 on U.S. 601, Ahles 12611. Marlboro Co.: 
Roadside, 3.6 mi. ne. of Marlboro-Chesterfield Co. line (Pee Dee River) 
on S. C. 77, Ahles 12613. Oconee Co.: Common on roadside, 1.5 mi. ne. 
of Fair Play on S. C. 59, Ahles 13360; roadside, 1.5 mi. ne. of Tugaloo 
River on S. C. 59, Bell 3085. Orangeburg Co.: Roadside, 1.1 mi. ne. of 
Orangeburg-Bamberg Co. line (South Edisto River) on U. S. 301, Ahles 
12607. Richland Co.: Roadside associated with Plantago aristata Michx., 
2.1 mi. n. of Calhoun-Richland Co. line (Congaree River on U.S. 601, 
Ahles 12609. 

In a revision of the Plantago patagonica complex Poe (1928) made the 
combination used here. "These plants, now established locally along the 
roads of North and South Carolina, are described by Small (1913) under 
the name Plantago Wrightiana Decne. as native to Texas. Kearney and 
Peebles (1951) give the range of this western plant as “western Texas 
to central Arizona, Oregon and California." It bas not been reported 
previously from the eastern United States. 

Sphenoclea zeylandica Gaertn.—sovTH CAROLINA. Jasper Co.: Weed 
in rice fields, Savannah River Wildlife Refuge, 1.3 mi. w. of jet. U. S. 17 
& 17A, and n. on levee, Ahles 18175. 

The range of this species is as yet too far south to be ineluded bv 
Fernald (1950) and Gleason (1952). Small (1033) gives the range as 


1958] Ahles, et al.,—Species New to the Carolinas 25 


*La. to Ark." Fox & Godfrey (1949) cite a specimen from Hyde County, 
North Carolina. In Jasper County it was found in only one rice field, 
but there fairly abundantly. 

Wahlenbergia gracilis Schrad.—sovTH CAROLINA. Aiken Co.: Weedy 
yard, about 2 mi. n. of North Augusta, H. Laing 174. Chesterfield Co.: 
Roadside, Cheraw St. Pk. near U. S. 52, Radford 15890. Colleton Co.: 
Weedy area along Seaboard Railroad on Co. Rt. 26, 8 mi. se. of U. S. 17, 
Ahles 12226. Darlington Co.: Damp soil back of Carnes farm about 5 
mi. s. of Hartsville, B. E. Smith 414; damp soil along highway between 
Hartsville and Darlington, B. E. Smith 415; damp flats near Lauther's 
Lake, B. E. Smith 416; damp soil on Witherspoon Island, B. E. Smith 
1730. Hampton Co.: Sandy field, 2.4 mi. ne. of Hampton on U. S. 601, 
Bell 3935. Lee Co.: Sandy roadside, 1 mi. sw. of Alcot on U. S. 15, Ahles 
15897. Marlboro Co.: Field, 2.5 mi. w. of Blenheim, Radford 12516; 
roadside, 1.5 mi. e. of Wallace near N. C. 90, Radford 15612; roadside 
Muddy Creek se. of Marlboro, Radford 19070. 

This species which is not found in the manuals by Small (1933), 
Fernald (1950) or Gleason (1952), has become very well established in 
many places in South Carolina and northern Florida. "The first collec- 
tion from South Carolina in the herbaria 1s dated 1939, from Darlington 
County. Extensive collecting in 1956 has yielded records from six addi- 
tional counties in this state. It is difficult to determine how long the 
species has been established in this region. 

Artemisia vulgaris ],—souTH CAROLINA. Laurens Co.: U. S. 221 at 
the Enoree River, Freeman 56920. 

Small (1933) in his range “Ga. to Ala., B. C., Ont. and Newf." would 
include this species as occurring in South Carolina, and Cronquist 
(1952) with the vague “established throughout most of eastern U. 8. 
and adjacent Canada," might also include South Carolina. Fernald 
(1950) gives the range “Nfld. to Ont., s. to N. S, N. E, Del, Pa. 
casually to Ga., Mich., Wisc. and Minn." Since there were no South 
Carolina specimens at NCU, DUKE or Ncsc, it seems that the Laurens 
County collection should be cited as the first record of the species for 
that state. 

Bidens pilosa L. var. radiata Bip.—souTH CAROLINA. Georgetown 
Co.: Sandy bank, causeway across marshes at Georgetown, Godfrey 
40911. Jasper Co.: Weed around Little City Motel, Ridgeland on U. 8. 
17, Ahles 12397. 

Sherff (1937) in his monographie treatment of the genus restricts the 
range of Bidens pilosa L. to the extreme southwestern United States 
and southward through Mexico. The plant in the eastern United States 
he refers to as var. radiata, but cites no specimens from South Carolina. 
Fernald (1950) says for this plant, “casual weed n. to Mass," and 
Cronquist (1952), “ballast waif at Philadelphia."  Sherff (1937) cites 
specimens from Massachusetts and New Jersey, indicating that it is 
sporadic at least as far north as Massachusetts. Wiegand (1933) gives 


26 Rhodora [Vor. 60 


the range as Florida, southern Georgia and Alabama. Godfrey (1950) 
cites one specimen from North Carolina. 

Chrysanthemum lacustre Brot.—NonTH CAROLINA. Chowan Co.: 
In ditch on N. C. 32, 1.9 mi. n. of Edenton, July 12, 1949, E. T. Browne, 
Jr. Stanly Co.: Low depression, 0.8 mi. nw. of Richfield, Ahles 16174. 

This species, native to Portugal according to Bailey (1950), is not 
included as an introduced plant in North America by any of the works 
cited in this paper. A specimen at DUKE labeled “Beach of Lake Huron, 
Mackinaw Island, Michigan, July 28, 1935, H. A. Gleason,” would seem 
to warrant its inclusion in the manuals of the northeastern United States. 
It is occasionally cultivated and apparently spreads from this source. 
The two specimens cited above constitute the first report for North 
Carolina, 

Cirsium arvense (L.) Scop.—NorTH CAROLINA. Ashe Co.: Flat bot- 
tomlands along New River, Sept. 21, 1946 (collector not given, Ncsc). 
Haywood Co.: Open grassy pasture, Crabtree Bald knoll, se. of summit. 
Plentiful in small area, Quarterman 85 (DUKE). Jackson Co.: Pisgah 
Forest, July 25, 1940, H. Myres. Swain Co.: Edge of abandoned road, 
s. of Black Camp, Blomquist 9757 (DUKE). Transylvania Co.: Open 
pasture in the Pink Beds, Correll 3389 (DUKE). 

Although Small (1933) gives the range as *N. C. to Kans., B. C. and 
Newf.," the preceding collections seem worthy of record, since Fernald 
(1950) gives the range south only to Virginia, while Cronquist (1952) 
says northern United States and southern Canada. 

Coreopsis basalis (Dietr.) Blake—sourH CAROLINA. Beaufort Co.: 
Cinder bed along railroad, 0.4 mi. e. of Burton on U. 8. 21, Ahles 12347. 
Colleton Co.: Railroad at Green Pond, on 8. C. 303, Bell 2326. Hampton 
Co.: Very abundant along railroad, Early Branch on S. C. 28, Ahles 
12445. 

Watson (1933a), under Coreopsis drummondii (D. Don) T. & G., 
gives the range of this species as Texas, and states that it is an escape 
from cultivation eastward. Fernald (1950) says that it spreads to waste 
ground. Cronquist (1952) reports it as found occasionally as an escape 
from cultivation. Godfrey (1950) reports it as abundantly naturalized 
around Southport, Brunswick County, North Carolina. In the three 
stations cited above for South Carolina, plants of this species were well 
established in good-sized stands, always near or on railroad ballast. 

Cosmos bipinnatus Cav.—souTH CAROLINA. Chesterfield Co.: Road- 
side, 2.5 mi. nw. of Jefferson, Radford 18792. 

Alexander (1933) gives the range of this species as Florida, Fernald 
(1950) says that it is becoming established southward, and Cronquist 
(1952) gives the range as Arizona and Central Mexico, cultivated else- 
where and frequently escaped. Thorne (1951) cites it from Georgia as 
infrequent along roadsides. No record of the occurrence of this species 
in South Carolina prior to 1956 has been published. 

Crepis pulchra [,—souTH CAROLINA. Chesterfield Co.: Waste place, 
Forked Creek, 1 mi. e. of Jefferson, Radford 12385. Oconee Co.: Once- 


1958] Ahles, et al.,—Species New to the Carolinas 27 


cultivated field, about 7 mi. s. of Westminster on Co. Rt. 20 at jet. Co. 
Rt. 140, Ahles 14047. Pickens Co.: Field, 0.7 mi. nw. of Norris on 8S. C. 
137, Bell 3458. Union Co.: U. S. 176, 4 mi. n. of Union, Freeman 56147. 

This taxon is not in Small (1933). Fernald (1950) gives the range 
as Virginia and Indiana. Cronquist (1952) gives it as Virginia, Ohio and 
Indiana. Fox & Godfrey (1949) report it from three counties in North 
Carolina, and as abundant at each locality. Freeman (1955) reports it 
as occasional in Polk County, North Carolina. During 1956 many addi- 
tional collections of it were made in North Carolina. The Oconee, 
Pickens and Union County collections in 1956 constitute the first report 
of this species from South Carolina. 

Facelis retusa (Lam.) Sch.-Bip.—sovTH CAROLINA. Allendale Co.: 
Sandy roadside, 0.1 mi. nne. of Co. Rt. 26 on Co. Rt. 49, Ahles 12523. 
Anderson Co.: Sandy roadside, 3 mi. ese. of Fair Play, Ahles 13416. 
Beaufort Co.: Cinder bed along railroad at station, Sheldon, Ahles 
10367. Colleton Co.: Weed in once-cultivated field, Ritter, Ahles 10538 ; 
sandy roadside, near Ashepoo River, about 1 mi. s. of Ritter on Co. Rt. 
41, Ahles 12189. Hampton Co.: Low woodland, 2.8 mi. se. of Furman 
on U.S. 621, Bell 1800. Jasper Co.: Lawn weed, Ridgeland, Ahles 10358 ; 
roadside, 5 mi. s. of S. C. 128 on S. C. 170, Ahles 12392. Newberry Co.: 
Indian Creek, 7 mi. sw. of Whitmire, Freeman 56223. Orangeburg Co.: 
Roadside near Bowman, common in waste places and meadows, Freeman 
56168. Pickens Co.: Dry upland pasture, 1.4 mi. w. of Pumpkintown, 
on S. C. 11, Ahles 14366. 

Under the binomial Facelis apiculata Cass., Small (1933) states the 
range as “coastal plain & piedmont, Fla., Ala. and Ga.” Fernald (1950) 
and Cronquist (1952) do not include the species. Thorne (1954) reports 
it as common in Georgia, and Godfrey (1950) cites a specimen from 
Cleveland County, North Carolina. Apparently it has become more 
abundant in the last few years and there are now specimens from many 
more North Carolina counties. In 1956 collections were made in nine 
counties of South Carolina, a state from which the species had not been 
reported previously. 

Helianthus maximiliani Schrad—NortH CAROLINA. Cleveland Co.: 
Open field along stream, 2 mi. ssw. of Mooresboro, Ahles 19200. Forsyth 
Co.: Open place (without further data), Schallert 610 (det. C. B. Heiser, 
1955). Rockingham Co.: Roadside, 1.75 mi. nw. of Lawsonville, Rad- 
ford 18569. 

Watson (1933), Fernald (1950) and Cronquist (1952) all give the 
range of this species as west of the Mississippi River, and state further 
that it is adventive, escaped or introduced eastward. 

Helianthus radula (Pursh) T. & G.—sovrH canorniNa. Colleton Co.: 
Sandy roadside of Co. Rt. 24, by pine woods, 0.5 mi. nw. of jet. with Co. 
Rt. 34, frequent locally, Bell 5382. Hampton Co.: Sandy upland, 2.2 mi. 
s. of Brighton on dirt road, Bell 4985; pine savannah, 0.4 mi. n. of 
Jasper-Hampton Co. line on S. C. 128, Ahles 18374; pine savannah, 0.1 
mi. w. of Hampton-Jasper Co. line on U. S. 321, 1 mi. e. of Garnett, 


28 Rhodora [Vor. 60 


Ahles 21009; pine savannah, 0.7 mi. ne. of Early Branch, Ahles 21019. 
Jasper Co.: Pine savannah, 2 mi. n. of Hardeeville on U. S. 321, Ahles 
18191; sand hill by Co. Rt. 41, w. of Barton and 0.5 mi. s. of jet. with 
Co. Rt. 23, Bell 5114; pine savannah and roadside, 3.3 mi. n. of Hardee- 
ville on U. S. 601, Ahles 20972. 

It is very surprising to note that Watson (1933) does not include 
South Carolina in the range of this species (“Fla. to Ala. and Ga."). 
In South Carolina it appears to be a very definite part of the flat pine- 
savannah flora. At each locality cited it was conspicuous and at least 
frequent, if not common. It was observed in many more localities 
within the cited counties, but specimens were not colleeted from all 
of them. 

Hypochaeris glabra L.—NwonTH CAROLINA. Alexander Co.: Open 
stream bank, Middle Fork of Little River near jet. with Catawba River 
sw. of Taylorsville, Radford 18167. 

No collections of this species were found from North Carolina at 
NCU, DUKE, Ncsc or Ny. Fernald (1950) does not include this species, 
while Cronquist (1952) says, “established in the Pacific states, and has 
been collected several times apparently as a waif in our range." Small 
(1933) gives a very ambiguous range, *n. Fla. to Ohio and Me." It 
is well established at the locality cited above and is here recorded as 
new for that state. 

Iva ciliata Willd.—NonTH CAROLINA. Anson Co.: Abundant on road- 
side and field edges for a stretch of one mile, 1 mi. s. of Pee Dee River 
on N. C. 109 and 1 mi. e. on dirt road, vicinity of Cedar Creek, Ahles 
19498. Franklin Co.: Weed in cultivated field, 0.4 mi. nw. of Sutton, 
Ahles 20664. Halifax Co.: Abundant on edge of cultivated field, 4.1 
mi. se. of Weldon on U. S. 301, Ahles 20820. 

The range of this species as a native plant is predominantly central 
United States and introduced eastward to Alabama according to Small 
(1933), and New England according to Fernald (1950). The three 
collections cited are all close to the fall line, two in the northern part, 
and one in the southern part of the state. Only at the Franklin County 
locality was this species scarce. 

Lapsana communis L.—xNonTH CAROLINA. Haywood Co.: Open moist 
hillside, Waynesville, lower part of trail to summit (of) Eaglenest, 
Quarterman 74 (DUKE). 

Fernald (1950) gives the range of this species as “Que. and Ont., s. 
to N. S, N. E, Va, W. Va. and Mo." Cronquist (1952) also has 
Virginia as the southernmost state on the Atlantie seaboard. Small 
(1933) does not include the genus. The above cited specimen is there- 
fore taken as a southward extension of the known range. 

Leontodon leysseri (Wallr.) G. Beck—NorTH CAROLINA. Clay Co.: 
Roadside, Buck Creek area near U. S. 64, w. of Black Gap, Radford 
16202. 

This species is not included by Small (1933). Fernald (1950) gives 
the range south to Ohio and New Jersey, while Cronquist (1952) has it 


1958] Ahles, et al..—Species New to the Carolinas 29 


south to the District of Columbia. The Clay County collection repre- 
sents a southward extension of the range. Superficially this species 
very closely resembles Hypochaeris glabra. 

Picris hieracioides L.—NonTH CAROLINA. Buncombe Co.: Habitat in 
vicinis Asheville, Maio, 1888, Gerald McCarthy. Haywood Co.: Juna- 
luska Mt., Blomquist 5291 (DUKE); moist open pasture, Waynesville, 
near summit of Eaglenest, Quarterman 73 (DUKE). 

Small (1933) does not include this species in his manual. Fernald 
(1950) says it occurs south to New Jersey and Pennsylvania, and Cron- 
quist (1952) south to the District of Columbia. 

Rudbeckia mollis Ell .—sovTH CAROLINA. Jasper Co.: Open sand 
hills, 8.5 mi. nw. of Tillman, 2.6 mi. se. of Seaboard Air Line Railway 
(this is presumably the town of Myres), Ahles 18216. 

This species has been reported by Boynton (1933) from Florida and 
eastern Georgia. The Jasper County specimen is therefore a northward 
extension of range. It is very probable that this is a native plant here, 
occurring as it does on the same sand ridge with Chrysobalanus oblongi- 
folius and Dicerandra densiflora. 

Soliva sessilis R, & P—NORTH CAROLINA. Carteret Co.: Lawn from 
Beaufort, May 1, 1953, A. H. Newsome (Ncsc). SOUTH CAROLINA. 
Orangeburg Co.: Weed in lawn, Edisto Gardens along Edisto River, 
Ahles 10695; same locality, Freeman 56165. 

Small (1933) gives the range of this species as “n, Fla. to La.” Thorne 
(1951) supplements this by reporting it from Georgia. The three above 
collections extend the range northward into South and North Carolina. 

Tragopogon dubius Scop—NortH CAROLINA. Catawba Co.: Weed 
along railroad, 2.6 mi. e. of Conover, Ahles 12984. 

Small (1933) does not include this species in his manual. Fernald 
(1950) gives it south only to Virginia, while Cronquist (1952) says, 
“established over most of the U. S., more common westward.” The 
absence of North Carolina specimens at NCU, DUKE, NCSC and NY seems 
to justify recording the species as new to North Carolina. 

Verbesina encelioides (Cav.) B. & H—NorrH CAROLINA. Robeson 
Co.: Scattered in pasture on upland, 20 mi. w. of Lumberton, Sept. 4, 
1953, B. W. Wells (Ncsc). SOUTH CAROLINA. Sumter Co.: Sandy waste 
ground, abundant, southern city limits of Sumter, AAles 15896 ; Wedge- 
field at the jet. of S. C. 763 & 261, Freeman 56804. 

Small (1933), under the name Ximenesia encelioides Cav., places this 
species as indigenous west of the Mississippi, and cites it as introduced 
in Alabama and Key West, Florida. Fernald (1950) has it adventive in 
Missouri and occurring casually to New England. Cronquist (1952) says 
it is occasionally introduced eastward. In the vicinity of Sumter, in 
Sumter County, South Carolina, this species is so abundant as to ereate 
large yellow masses in almost every vacant lot. 


30 Rhodora | Vor. 60 


SUMMARY 


One hundred state records are reported: fifty taxa are reported 
for the first time from North Carolina, and fifty taxa from South 
Carolina. Two hybrids and only four subspecific taxa are included 
in the records, and of these latter, two varieties, Plantago hooker- 
iana var. nuda and Bidens pilosa var. radiata, also represent 
species new to the respective states for which they are reported. 
Of the total of eighty-eight taxa treated as new to one or both 
of the Carolinas, fourteen taxa represent native plants, sixty- 
seven taxa represent introduced plants, and seven taxa (Digitaria 
filiformia, D. texana, Paspalum plicatulum, Scirpus koilolepis, 
Phyllanthus pentaphyllus, Callitriche terrestris and Ampelamus 
albidus) represent plants of questionable origin in the Carolinas. 
Vouchers for all records are deposited in the Herbarium of the 
University of North Carolina unless otherwise indicated. 


LITERATURE CITED 


ALEXANDER, E. J. 1933. Cosmos, in J. K. Small, Manual of the South- 
eastern Flora, p. 1454. 

Baitey, L. H. 1950. The Standard Cyclopedia of Horticulture, 3 vols. 
2nd ed. MacMillan, N. Y. 

BkETLE, A. A. 1947. Scirpus, in North American Flora 18: 479—504. 

Boynton, C. L. 1933. Rudbeckia, in J. K. Small, Manual of the South- 
eastern Flora, pp. 1422-1427. 

CHasE, AGNES. 1950. Manual of the Grasses of the United States, United 
States Govt. Printing Office, Washington, D. C. 

Cronquist, A. J. 1952. Compositae, in H. A. Gleason, Illustrated Flora 
of Northeastern United States. 3: 323-545. 

Curtis, M. A. 1867. Geological and Natural History Survey of North 
Carolina. Botany, 3: 1-156. Raleigh, N. C. 

Duncan, Winpug H. 1950. Preliminary reports on the flora of Georgia—4. 
Notes on the distribution of flowering plants including species new to 
the state. Castanea 15: 146-159. 

Fassert, Norman C. 1951. Callitriche in the New World. Ruopora 53: 
137-155, 161—182, 185—194, 209-222. 

Fernatp, M. L. 1938. Noteworthy plants of southeastern Virginia, RHno- 
DORA 40: 364—424, 434—459, 467—485. 

——————— 1941. Another century of additions to the flora of Virginia. 
Ruopora 43: 485—553, 559-630, 635—657. 

1950. Gray's Manual of Botany, ed. 8, 1632 pp., American 
Book Co., N. Y. 

Fox, W. B. & R. K. Goprrey. 1949. Notes on distribution of North Caro- 
lina plants—I. Ruopora 51: 129-146. 

FREEMAN, OtiveR M. 1955. Notes on the flora of Polk County, North 
Carolina. Castanea 20: 37-57. 


1958] Ahles, et al.,—Species New to the Carolinas 31 


GLEASON, H. A. 1952. Illustrated Flora of Northeastern United States. 
3 vols. New York Botanical Garden, N. Y. 

Goprrey, R. K. 1950. Studies in the Compositae of North Carolina III. 
An enumeration of noteworthy distribution records. Jour. Elisha 
Mitchell Sci. Soc. 66: 186-194. 

Heci, G. 1912. Illustrierte Flora von Mittel-Europa. Vol. 3, Dicoty- 
ledones, pt. 1. Munchen. 

Hitcucock, A. S. 1933. Poaceae, in J. K. Small, Manual of the South- 
eastern Flora, pp. 30-139. 

Hoover, R. F. 1941. A systematic stuty of Triteleia. Am. Midl. Nat. 
25: 73-100. 

JAMES, Ropert L. 1956. Introduced plants in northeast Tennessee. Cas- 
tanea 21: 44-52. 

Kearney, T. H. « R. H. Prestes. 1951. Arizona Flora. 1032 pp. Univ. 
of Calif. Press. Berkeley and Los Angeles. 

Macsripe, J. Francis. 1930. Spermatophytes, mostly Peruvian—II. Field 
Mus. Bot. Ser. 8: 77-130. 

Macue, BassETT. 1952. Caryophyllaceae, in H. A. Gleason, Illustrated 
Flora of Northeastern United States. 2: 118-145. 

Martuias, MILDRED & Lincoun Constance. 1944-45. Umbelliferae, in 
North American Flora 28B: 43-295. 

MorpENKE, H. N. 1949. The known geographic distribution of the mem- 
bers of the Verbenaceae, Avicenniaceae, Stilbaceae, Symphoremaceae 
Eriocaulaceae. 215 pp. Privately published. New York. 

1952. Verbenaceae, in H. A. Gleason, Illustrated Flora of 
Northeastern United States. 3: 125-139. 

Musz, Pniur A. 1938. Studies in Onagraceae XI. A revision of the 
genus Gaura. Bull. Torrey Club 65: 105-122, 211-228. 

PENNELL, F. W. 1933. Rhinanthaceae, in J. K. Small, Manual of the 
Southeastern Flora, pp. 1182-1225. 

Por, Ione. 1928. A revision of the Plantago patagonica group of the 
United States and Canada. Bull. Torrey Club 55: 406-420. 

Rankin, H. A. 1940. Vacation notes. Jour. Gray Memorial Bot. Assoc. 
8: 10-11 (Mimeographed). 

Ropars, C. L. 1950. The Umbelliferae of North Carolina and their 
distribution in the Southeast. Jour. Elisha Mitchell Sci. Soc. 66: 
195-266. 

Rocers, Davin James. 1951. A revision of Stillingia in the New World. 
Ann. Mo. Bot. Gard. 38: 207-259. 

SuerFrr, E. E. 1937. The genus Bidens, Part I Publ. Field Mus. Nat. 
Hist. Bot. Ser. 16: 1-709. 

SMALL, J. K. 1895. A monograph of the North American species of the 
genus Polygonum. Mem. Dept. Bot. Columbia College. 1: 1-183. 
——— 1913. Flora of the Southeastern United States. Published 

by the author. N. Y. 
- 1933. Manual of the Southeastern Flora. Published by the 
author. N. Y. 

SMirH, Bupp E. 1946. Additions to the flora of South Carolina. Jour. 
Elisha Mitchell Sci. Soc. 62: 81-86. 

THonwNE, RoBERT F. 1951. Vascular plants previously unreported from 
Georgia. Castanea 16: 29-48. 


32 Rhodora [ Vor. 60 


1954. The vascular plants of Southwestern Georgia. Am. 
Midl. Nat. 52: 257-327. 
WALTER, THOMAS. 1788. Flora Caroliniana, 263 pp. London. 
WarsoN, E. E. 1933. Helianthus, in J. K. Small, Manual of the South- 
eastern Flora, pp. 1431-1441. 
1933a. Coreopsis, in J. IK. Small, Manual of the South- 
eastern Flora, pp. 1446-1450. 
WiEGAND, K. M. 1933. Bidens, in J. K. Small, Manual of the South- 
eastern Flora, pp. 1450-1454. 


BanrHoLoMEW's CoBBLE.—This unpaged pamphlet briefly describing 
the geological and botanical features of a rocky 30 acre reservation near 
Ashley Falls, Massachusetts, is noteworthy because of the accurate 
list of plants of the area provided. The pamphlet is made attractive 
with six full-page photographs of a few of the interesting plants found 
at the Cobble and of the Cobble itself. The plant list is mostly based 
upon studies made by the late C. A. Weatherby, who was extremely 
interested in having the natural features of the area preserved for the 
enjoyment of present and future generations. It is fortunate that the 
Cobble is in the hands of the Trustees of Reservations so that preser- 
vation is now assured. The Trustees of Reservations is a non profit 
corporation composed of publie-spirited citizens in Massachusetts who 
have an interest in preserving distinctive parts of our heritage from 
threatened destruction or irreparable alteration. The organization has 
been in existence for over 65 years and has by acquisition and manage- 
ment done much to make places of beauty and of historical interest 
within the Commonwealth available to the public.—R. C. Rous. 


1 Bartholomew’s Cobble, by Herbert J. Arnold and S. Waldo Bailey. Published by, and 
available from the Trustees of Reservations, 50 Beacon Street, Boston 8, Mass., $0.50. 


Volume 59, number 708, including pages 293—321 was issued 21 January, 
1058. 


79 
K 4? 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS l 
RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR, ( 


IVAN MACKENZIE LAMB 


Vol. 60 February, 1958 No. 710 

CONTENTS: 

Distribution of Littorella americana in the Mid-arrowed Region of 

Minnesota. Olga Lakea o. oe ene LI e is 33 
Experiments and Observations Bearing on Evolution in Oenothera. 

B NRINNN Gales errr E T a ie 37 
Plant Notes from Illinois. Glen J. Winterringer..............-. 41 
Arenaria rossii and Some of Its Relatives in America. 

Bassett Maguire... sees so sce ce cy ee ae a EET 
Rumex stenophyllus in North America. Doris Lóve and Frére 

Jean-Paul Bernard ........:. 4 de DETUR 54 
Elatine triandra in New York. Joseph Monachino.............. 58 
Species Plantarum—More Than a Facsimile Edition. 

R: C. Rollins (Review) |... eee ee Eee ere 59 


The New England Botanical Club, Inc. 


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CARD-INDEX OF NEW GENERA, SPECIES AND 
VARIETIES OF AMERICAN PLANTS 


For all students of American Plants the Gray Herbarium Card-index 
of Botanical Names is indispensable. It is a work of reference essen- 
tial to scientific libraries and academies and all centers of botanical 
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Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 February, 1958 No. 710 


DISTRIBUTION OF LITTORELLA AMERICANA IN THE 
MID-ARROWHEAD REGION OF MINNESOTA 


OrcA LAKELA 


Professor Fernald in separating L. americana from L. uniflora 
(L.) Asch.’ refers to the plant as one of the rarest in North 
Ameriea, known only from a few localities throughout its range 
from Newfoundland to Minnesota. Until recently the collection 
of L. H. Bailey, no. 437, Basswood Lake, July 28, 1886 appears 
to have been the only record from the state. 

L. americana first came to my attention while I was collecting 
in the “Roadless Area" of St. Louis County. Plants without 
flowers or fruit (no. 16743, August 9, 1953) were collected from 
a submersed colony with Lobelia Dortmanna, growing on a sandy 
bottom in shallow shorewaters of Iron Lake on the Ontario border. 
On the following day it was found again in Lac La Croix, at 
Beatty portage from Loon Lake. Plants in vegetative condition, 
no. 16756, were collected from a colony submersed in shallow 
water, again associated with L. Dortmanna. In each site the 
bottom soil was mostly a mixture of gray sand with black organic 
soll, peaty or mucky. Identification of the species in vegetative 
condition remained doubtful until 1957. In dealing with the 
known flora from Lake County for the manuscript of a Flora 
of the Mid-Arrowhead Region, Bailey's early collection of flower- 
ing plants was studied. This decisively cleared the identity of 
the sterile specimens. 

In the ensuing search for additional flowering material in late 
season Basswood Lake seemed most accessible. Working from 


1 Ruopora 20: 61-62. 1918. The North American Littorella. 


34 Rhodora [Vor. 60 


the Quetico-Superior Wilderness Research Center at Basswood 
Lake, after an extensive fruitless search of the more distant 
shores, Littorella was sighted without effort in the “home harbor” 
shore of the Center bay near the boat docks. The compact 
colony growing in silty sand was stranded above the water level. 
The associate species was Ranunculus repens. Coll. no. 22417, 
Sept. 10, 1957, consists of plants in late anthesis, with some 
mature fruits. 

The label data of Bailey’s collection lacks a specific location 
on Basswood Lake. The finding of the species there again tempt- 
ingly invites one to visualize Dr. Bailey towering over the spot 
preferred by a persistent colony of Littorella. However, in re- 
ality, the occurrence of a solitary colony on a lake with several 
hundreds of miles shoreline is presumptuous. 

L. americana was encountered again in Snow Bank Lake, lo- 
cated in the north central part of Lake County, about three miles 
south of the Canadian border. Here, in peaty sand of shallow 
waters of the bay south of the Resort, Subularia aquatica, in 
late flower and fruit, occurred in abundance with Isoetes muri- 
cata. Farther south in an adjoining bay some dozen plants of 
Nymphaea tuberosa were in full flower. Not far from the latter 
site on the sandy shore was an emersed colony of sterile Littorella, 
no. 22492 Sept. 14, 1957. Perhaps it was the stranded part of a 
much larger submersed colony 5-10 feet from the beach at a 
water-depth of 2-3 feet which covered square yards of the sandy, 
cobblestone strewn bottom. The plants were clearly visible in 
the early morning sun, but almost beyond the reach of the col- 
lecting tool. Only a few plants dislodged from the seemingly 
hardened sediment floated to the surface with fragments of 
Myriophyllum tenellum. 

It may be permissible to state here that botanizing is catching. 
On finding Littorella at the Quetico-Superior Research Center, 
the plants were shown to Mr. Otto Oltman, foreman, with a re- 
quest that he try to find and collect the species during an antici- 
pated canoe trip through the wilderness canoe country. His 
collection of Littorella, from the shore of an island in Malberg 
Lake about 4 miles west of Cook County, Sec. 8, T. 68, R. 6 W, 
September 19, 1957, was a welcome contribution to knowledge 
of this little known species. Thus within a space of a week, 


1958] Lakela,— Distribution of Littorella americana 35 


three new localities were discovered. The Ontario site in the 
accompanying map 1s based on a sight record made September 
2, 1956; circumstances prevented collecting at the time. "The 
colony may be found on the shore of a small bay connected 
with Crooked Lake by a narrow channel, opposite Curtain Falls 
Resort Area. 


4 
- 
= 
A 
> 
, 
- 
le 


9» 


95 


Ap 
b 


LirioRELLA AMERICANA. Fig. 1, at left, sheathing leaf-base, X 30; tissues stippled, clear 
areas air-chambers or lacunae, vascular traces cross-hatched. 

Fig. 2, at right, cross section of leaf near the middle, X 50; cells in outline only; epi- 
dermis without chlorophyll; (elongate cells with straight walls in face view not shown); 
stomata numerous throughout; mesophyll spongy with radial lacunae; traces cross-hatched. 


In studying living plants of Littorella discrepancies in descrip- 
tions of leaves by different authors came to my attention. Ac- 
cording to N. C. Fassett,? the leaves are “rather stiff dark thread- 
like." H. A. Gleason? notes their shape as “linear.” In his illus- 
tration of the plant as a whole, they are depicted as being flat 
and thin. Professor Fernald features leaf morphology as one of 
the diagnostic differences between the American and European 

2 Manual of Aquatic Plants p. 313-314, 1940. 


3 The New Britton and Brown Illustrated Flora of the United States and Canada Vol. 3, 
p. 273, 1952. 


36 Rhodora "Vor. 60 


species. In describing L. umericana, he observed the leaves as 
“flattish, faleate-arcuate or straightish"; in L. uniflora, as “sub- 
terete or semi-cylindric." 

It may not be amiss to place on record another description 
based on the study of living plants from five different localities. 
Mature fully turgid leaves are subulate, faleate-arcuate, lustrous, 


ELA 


ST. 


Fig. 3. The known sites of Littorella americana, in the upper portions of St. Louis and 
Lake Counties, mid-Arrowhead Region, Northeastern Minnesota. 


bright green to yellowish green above the white bases. The blade 
distally above the shallow groove of the sheathing leaf-base, feels 
and looks terete, firm and pliable. The rich green tips of young 
leaves, two in alternate succession, embraced within the searious- 
margined bases of opposing mature leaves are somewhat com- 
pressed but soon become subulate. Microscopically the mature 
leaves are nearly terete or at least more than semi-circular with 
concentric mesophyll centered about the median trace. The large 
air chambers appear to be radial; the two smaller traces are 
elevated above the median plane, ef. fig. 1 & 2. 


1958] Gates, —E volution in Oenothera 37 


Although Littorella uniflora, the European species has not been 
studied, descriptions of its leaves as, “subterete or semi-cylindric" 
indicates a similarity to those of L. americana. Otherwise, in 
floral struetures and size Minesota plants well agree with Fern- 
ald's descriptions. The purplish-black fruit in maturity appears 
terete, apieulate with a short stipe and a minutely rugose pericarp. 

Plants collected in late October show yellowing and gradual 
decay of the older leaves. Under greenhouse conditions the young 
leaves continue growth. The renewed overwintering rhizome of 
the season is 2-3 mm. thick and about as long; rhizomes of the 
previous years are persistent, subject to gradual decay. Whether 
the plants are stranded or submersed, they are readily recognized 
in field studies. The terete-appearing leaves, 1-2.2 mm. thick 
near the midpoint cannot be confused with Ranunculus repens.” 
Their outwardly-arching habit sets them apart from the linear- 
compressed obtuse leaves of Lobelia Dortmanna, which are 
broadly elliptic in cross-section, with two lacunae flanking the 
median trace. 

The author is indebted to the Quetico-Superior Wilderness 
Research Center for courtesies pertinent to facilities for field 
studies, the Graduate School of University of Minnesota for 
defraying the cost of collecting and Dr. J. B. Carlson and Donald 
W. Davidson, Duluth Branch, for preparing the illustrations.— 
UNIVERSITY OF MINNESOTA, DULUTH BRANCH, 


EXPERIMENTS AND OBSERVATIONS BEARING ON 
EVOLUTION IN OENOTHERA 


R. RUGGLES Gates? 


I 


During an examination of the collections of Oenothera in the 
Gray Herbarium, Harvard University (Gates, 1957), a new 
species Oe. perangusta (Gates, 1950) was described from the 
North shore of Lake Superior. One specimen in the collection 
from Jackfish Station differed from the rest in having deep red 
stems and buds. It was recognized as a mutation parallel to 
the red-budded mutation from Oe. Lamarckiana (Gates, 1911) 


1 Cambridge, Mass. 


38 Rhodora [Vor. 60 


and was therefore called Oe. perangusta var. rubricalyx. While 
in Vancouver, B. C., in 1953 I found in the herbarium of the 
University of British Columbia, through the courtesy of Professor 
Hutchinson, specimens of the same species and its red-budded 
variety originally collected from the same locality. On my way 
East I was able to stop at Jackfish, and had the good fortune 
to find the original locality from which the red-budded mutation 
was derived. 

About 1400 feet east of Jackfish Station on August 28, 1953 
near the railway, a colony of Oenothera was found in which were 
counted 7 plants with red stalks in fruit and many young rosettes 
also evidently of the red variety (as shown by the red colour 
ventrally of the midribs), as well as one plant of the ordinary 
type with green stems. A specimen was collected for the Gray 
Herbarium. The red plants in this clump must all have been 
descended from the original mutation. How old this colony is 
can only be conjectured, but Mr. Peter Leschuk, who managed 
the local hotel and afterwards sent me seeds on Sept. 28 when 
they were ripe, thought he remembered seeing the red form 
here as a boy. The clump might easily be destroyed by railway 
operations. Search of the area failed to reveal more than the 
one group of red plants. 

In June, 1954, returning from Japan over the same route, many 
observations of Oe. perangusta were made. The species was very 
uniform all along the north coast of Lake Superior from Schreiber 
Station, where there were large colonies, to Terrance Bay, near 
Angler Station, at Marathon and along the C.P.R. line to Heron 
Bay. At the last locality were large numbers of plants in their 
favorite habitat, loose sand and gravel on the steep railway 
embankment some 400 yards west of the station. Nowhere were 
red plants seen except in the clump near Jackfish Station. The 
same species was afterwards seen growing by the railway near 
Hamilton, Ont. It thus evidently occupies a wide area in Ontario, 
from the north shore of Lake Superior to Hamilton on Lake 
Ontario and the Bruce Peninsula (Gates, 1950) of Lake Huron. 


II 


All the small-flowered Oenotheras, including nearly all the 
species in Canada, are self-pollinating. "There is clear evidence, 


1958] Gates,— Evolution in Oenothera 39 


however, that cross pollination occasionally occurs between dif- 
ferent forms occupying the same area. Such crosses are an im- 
portant factor in the evolution of the genus, and it is therefore 
desirable to obtain some evidence of the frequency with which 
cross pollination takes place in nature. For this purpose the ideal 
would be to use Oe. perangusta and its dominant red mutation, 
planting them in alternate rows and collecting open-pollinated 
seeds from the green form. Any plant with red buds derived 
from these seeds would then be the result of cross-pollination. 

Before seeds of Oe. perangusta and its red form were available, 
this experiment was tried with Oe. Victorini, a species with some- 
what larger flowers which may be somewhat less strictly self- 
pollinating. Seeds of Oe. Victorini were obtained from the Mon- 
treal Botanical Garden through the Director, Professor Jacques 
Rousseau, and of a strain of Oe. blandina containing the gene 
(rubricalyx) for red buds from Professor D. G. Catcheside. 
These were grown at the Bussey Institution, Jamaica Plain, 
Mass., the facilities being kindly provided by Professor Karl Sax 
of Harvard University. The two species were planted in four 
alternate rows, ten plants to each row. When the seeds were 
collected on September 15, 1953, only nine plants of Oe. blandina 
rubricalyx had flowered, mostly from side branches. They also 
came into flower later than Oe. Victorini. The 20 plants of this 
latter species all flowered and were full of seeds, many of the 
capsules having already shed some of their seeds. Since the 
species with red buds began flowering later, only the later seed 
capsules of Oe. Victorini were collected, four capsules from each 
of 12 plants. 

As a partial control of the frequency of crossing, the number 
of capsules on 12 plants of Oe. Victorini was roughly estimated. 
They totalled 2305, with a range from 100 to 385 per plant. 
Estimating 300 seeds per capsule, a total of 691,500 seeds could 
have been exposed to “red” pollen in time to ripen before the 
frosts. It was judged that 10 lower capsules per Victorini plant 
or 36,000 seeds, were fertilized before the blandina pollen began 
to be produced. Subtracting this number leaves 555,500 seeds 
probably exposed to “red” pollen. 

About four capsules each from eleven exposed Victorini plants 
were sown on vermiculite in the greenhouses of the Bussey Insti- 


40 Rhodora [Vor. 60 


tution on Ooctober 2, 1953. On January 16, 1954, they had 
produced 3395 young rosettes, two of which had ventrally red 
midribs and would therefore have red buds. "This gives à very 
tentative crossing frequency of 1:1698. Later attempts to get 
results on a large scale, using Oe. perangusta and its red variety, 
have not succeeded because the plants remained rosettes which 
failed to survive the winter season. 


HI 


One incidental observation is worth recording. Oenothera 
flowers are generally visited by nocturnal moths after the flowers 
open in the evening. Their long proboscis enables them to suck 
up the nectar which is secreted in the hypanthium and fills the 
lower part of this tube. Bees may also be seen visiting Oenothera 
flowers. One bee which was carefully observed visited flower 
after flower in a routine way. Being unable to obtain the nectar 
at the base of the hypanthium in the normal way by sucking 
it up from the inside, it lighted on a petal, then walked down 
the slender hypanthium, punctured it at the base just above the 
ovary, and lapped up the nectar, leaving an ooze of nectar where 
its short proboscis had been withdrawn. The pollination mechan- 
ism was thus entirely bypassed, the bee obtaining the nectar 
without entering the flower. This was done with flower after 
flower on different plants as a regular routine. Whether this 
bee was exceptional in having discovered a way to circumvent 
the floral mechanism, or whether this is a general custom of 
bees in the New England area or elsewhere is unknown. It 
shows at any rate that some bees have developed an efficient 
method of their own for extracting the nectar from Oenothera 
flowers. The fact that each flower is punctured in the position 
to obtain the maximum amount of nectar from the nearly erect 
hypanthium, seems to show a mental activity closely akin to 
intelligence. 

LITERATURE CITED 
Gates, R. R. 1911. Studies on the variability and heritability of pig- 
mentation in Oenothera. Zeits. f. Abst. u. Vererb. 4: 337-372. 
1950. Another parallel mutation in Oenothera. Canad. 
Field-Naturalist 64: 142-145. 


— 1957. A conspectus of the genus Oenothera in North 
America, RHODORA. 57: 9-17. 


1958] Winterringer,—Plant Notes from Illinois 41 
PLANT NOTES FROM ILLINOIS 
GLEN S. WINTERRINGER 


Several plant species of western United States have made their 
way into Illinois and have apparently become established, e.g., 
Froelichia gracilis (Hook.) Moq., Callirhoe involucrata (T. & 
G.) A. Gray, Salvia pitcheri Torr., Salvia reflera Hornem. and 
several others. The newest arrival is Phlox gracilis (Hook.) 
areene which according to William A. Weber is, “a common but 
inconspicuous weedy annual of the mesas and foothills”. Such 
a habitat sounds strange when applied to plants collected in 
Illinois for this little weed was found in mid-May growing along 
the shoulder of a main highway west of Niantic in Macon County. 
It seems likely to the writer that this small weedy annual may 
have grown in the area for some time and has been overlooked. 
It would be interesting to know how the plant actually arrived 
in the Illinois area. In these days of superhighways and trans- 
continental travel it is not surprising that seeds or even plants 
are transported from one corner of the continent to another. 
What may be more important is the ability of a species, even a 
weed, to adapt itself in this new locality. Continued observations 
of this weedy Phlox will be made during subsequent growing 
seasons. Collection data. Macon Co.: May 12, 1957, shoulder 
of Highway No. 36, 215 mi. W. of Niantic, G. S. Winterringer 
14003. Another collection same locality May 16, 1957, G. S. 
Winterringer 14001, 14002. Specimens are in herbaria of the 
following: Illinois State Museum, Illinois Natural History Sur- 
vey, and the University of Colorado. 

The following items are listed either (a) they are new for the 
state or (b) they are very infrequently collected plants in Illinois. 
'The writer expresses thanks to R. T. Rexroat of Virginia, Illinois, 
for his contribution of excellent botanical specimens. 


SPECIES NEW TO THE STATE 


SCLERIA RETICULARIS Michx. Cass Co.: east of Beardstown, Sept. 14, 
1956, in sand at edge of peaty sand hole, R. T. Rerroat 3438. Specimen 
verified by Earl T. Core. In August, 1957, Rexroat reported an 
abundance of this species in the same area. 

FaLcARIA srOIDES (Wibel) Aschers. Schuyler Co.: 2 mi. W. of 
Frederick, June 28, 1955, moist soil, R. T. Rexroat 1800. Verified by 


42 Rhodora [Vor. 60 


Mildred E. Mathias. "This species was not recently observed in the 
same area. 


INFREQUENTLY COLLECTED PLAN TS 


ARISTIDA DESMANTHA Trin. & Rupr. Morgan Co.: Oct. 12, 1956, E. of 
Meredosia; dry and sandy, R. T. Rexroat 3516, 8517. Cass Co.: Sept. 
14, 1956, E. of Beardstown; sand, R. T. Rexroat 3434, 3435, Mason 
Co.: Oct. 9, 1955, W. of Saidora, sand, R. T. Rexroat, 2426, 2403, 
2427, 2428. Oct. 18, 1954, with no definite county locality, dry sand, 
not common, R. T. Rexroat 1294. The Mason Co. specimens verified 
by Jason R. Swallen. Originally collected in Illinois by M. S. Bebb 
in 1861. Since that time it was collected by V. H. Chase in 1929 and 
by H. E. Ahles in 1950. The Morgan and Cass County locations are new. 

ECHINODORUS PARVULUS Engelm. Cass Co.: Aug. 13, 1957, E. of 
Beardstown, moist peaty sand, R. T. Rexroat 4150, 4151, 4152. The 
collector reported numerous plants of this species in the area. Previous 
Cass and Mason County records are indefinite. A collection was made 
in St. Clair County in 1892. 

ZIZANIA Aquatica L. Mason Co.: Aug. 19, 1956, S. of Havana; in 
shallow spring water, R. T. Rexroat 3225, 3226. The reason for report- 
ing these colleetions is the apparently unusual length of the eulms which, 
according to the collector, were from 11 to 13 ft. in length. Leaves, 
near the upper part of the culm, 2 in. wide and wider on the lower part. 

TRADESCANTIA BRACTEATA Small. Mason Co.: May 31, 1953, SW. part 
of the county, in sand and sun, fls. blue, R. T. Rexroat. 182, and same 
date, fls. rose, R. T. Rexroat 763. 

TRILLIUM RECURVATUM, forma SHAYI Palmer & Steyerm. Cass Co.: 
May 14, 1953, NE. of Virginia in woods, R. T. Rexroat 85. 

IRIS BREVICAULIS Raf. Morgan Co.: June 15, 1955, 10 mi. NW. of 
Jacksonville, wooded clay hillside, R. T. Rexroat, 2008. Cass Co.: June 
6, 1955, NE. of Virginia, elayey woods, R. T. Rexroat 2001. Same 
locality June 8, 1955, R. T. Rexroat 2002, 2003, 2004. Same locality 
July 22, 1955, R. T. Rexroat 2005, 2006, 2007 (with capsules). 

RuBUs LACINIATUS Willd. Mason Co.: July 14, 1957, E. of Bath, 
on a small sand dune with wild plum, e. T. Rexroat 3981, 3982. 

RUBUS PHOENICOLASIUS Maxim. Morgan Co.: May 31, 1956, 5 mi. 
SE. of Meredosia, R. T. Rerroat 2735. June 14, 1955, W. of Arenzville, 
R. T. Rexroat 1776, 1777. This introduced Rubus has been regarded 
previously as not established in Illinois. At the Arenzville locality the 
plants were abundant and thoroughly established over a considerable 
area, 

Lorus cornicutatus L. Cass Co.: June 4, 1953, sand, R. T. Rerroat 
201. July 17, 1953, fence row in full sun, R. T. Rexroat 389. Mason 
Co.: July 8, 1956, 5 mi. S. of Havana, dry, peaty sand, R. T. Rexroat 
2988. Brown Co.: August 30, 1956, N. of Cooperstown, clay hillside, 
R. T. Rexroat 3344. 


1958] Winterringer, —Plant Notes from Illinois 43 


CALLIRHOE INVOLUCRATA (T. & G.) A. Gray. Schuyler Co.: July 25, 
1956, N. of Browning, clay; roadside, stems trailing 2-3 ft, R. T. 
Rexroat 3099, 3100. 

ECHIUM VULGARE L. Schuyler Co.: June 28, 1955, 3 mi. S.W. of 
Frederick, clay hillside in pasture, A. T. Rexroat 1807, 1808, 1809. A 
few plants were observed in July 1957. 

TRICHOSTEMA DICHOTOMUM L. Cass Co.: Sept. 17, 1953, no definite 
locality, moist sand, R. T. Rexroat 684. Aug. 19, 1954, peaty sand, dry 
or moist, R. T. Rexroat 1358, 1102. Sept. 8, 1955, 4 mi. E. of Beards- 
town, in peaty sand, R. T. Rexroat 2285, 2286. In these Cass County 
localities this species was abundant. 

SALVIA PITCHERI Torr. Cass Co.: Sept. 11, 1954, near Virginia, dry 
soil, full sun, R. T. Rexroat 1359. Sept. 11, 1954, along R. R. tracks 
in full sun, perennial with several stems, R. T. Rexroat 1188. Sept. 
10, 1954, R. R. tracks in full sun, R. T. Rexroat 1189. Sept. 23, 1955, 
near Virginia, dry clay soil, R. T. Rexroat 2372, 2371, 2370. Plants 
of the areas cited above have been destroyed but specimens transplanted 
to Rexroat's garden were in good condition in July 1957. 

SALVIA REFLEXA Hornem. Woodford Co.: Aug. 6, 1957, 4 mi. S. of 
El Paso in an old pastured area, G. S. Winterringer 13818, 13819. The 
abundance of this adventive western sage in this locality was surprising. 
It was the dominant species over approximately an acre. 

MENTHA ALOPECUROIDES Hull. Schuyler Co.: July 25, 1956, N. of 
Browning, clay soil, spreads by stolons, R. T. Rexroat 3102, 3103. 
Specimens transplanted to Rexroat's garden in 1956 are now thriving. 

MiMvuLus GEYERI Torr. Mason Co.: May 13, 1956, 5 mi. S. of 
Havana, growing in a spring-fed creek in an inch of water, forming a 
dense mat, R. T. Rexroat 2612. 

All specimens cited in this paper are deposited in the Illinois State 
Museum Herbarium.—ILLINOIS STATE MUSEUM, SPRINGFIELD, ILLINOIS. 


44 Rhodora [Vor. 60 


ARENARIA ROSSII AND SOME OF ITS RELATIVES 
IN AMERICA 


BASSETT MAGUIRE 


Recently, my colleague, Arthur Cronquist, collected a series 
of an interesting Arenaria of the rossii-complex in Montana that 
seemed to require special study. About the same time, William 
A. Weber sent to me collections from Colorado which bore on a 
related problem and required further consideration of the status 
of Arenaria macrantha (Rydb.) Nels. (also of the A. rossii- 
complex). Further, a recently published paper by Nannfeldt! 
raised questions as to taxonomic interpretation and status, and 
the typification and nomenclature of Arenaria rossii. All of 
these circumstances made necessary a review of the matter and 
require the present statement. 

In his discursive paper, Nannfeldt has presented arguments 
purporting to establish the name (Minuartia) rolfii to supplant 
the long-used epithet rossii. Examination of historical materials 
at the British Museum (BM), Kew (x), Fielding Herbarium, Ox- 
ford (oxr), Gray Herbarium (GH), and The New York Botanical 
Garden (Ny), and a large body of newly collected specimens on 
deposit at the National Herbarium (cAN) and the herbarium of 
the Department of Agriculture (pao), both at Ottawa, the Gray 
Herbarium, and The New York Botanical Garden, has convinced 
us that there is but one polymorphic arctice American species in- 
volved, and that all of the Riehardson specimens and those of 
the Parry Voyage belong to it. Accordingly, there is no necessity, 
or indeed permissibility, for nomenclatural change or substitution, 
since it is clear that Richardson did in fact effectively and 
validly publish the epithet Rossii (Franklin Journ. p. 738. 1823), 
and that neither the epithet nor the authorship by Richardson 
may be displaced.” 

Arenaria rossii in the broad sense extends in the American 
Arctic from the Eastern Arctic Archipelago and northeast Green- 
land (also in Spitzbergen, ace. to Nannfeldt) westward to the 
Bering Straits and the Aleutian Archipelago, and in the moun- 


! Nannfeldt, J. A. Some Notes on Minuartia stricta (Sw.) Hiern. and Allied Species. 
Nytt. Mag. Bot. 3: 159-170. 1954. 

? This position was substantially taken by Porsild in Vase. Pl. West Cana. Arctic Archi- 
pelago Bull. 135, Nat. Mus. Cana. p. 110, 1955. 


1958] Maguire,—Arenaria rossii 45 


tains in Alaska and Yukon south to Wyoming and Oregon. 
Clinal modifications take place westerly by which the sepals be- 
come acute, the petals narrower and shorter, and the leaves less 
fleshy. The occidental population has been known as A. elegans. 
The line of phytogeographie division between the two races seems 
to lie just west of the delta of the Mackenzie River, perhaps to 
the north of the Richardson Mountains. 

In the Mackenzie Range and southward in the Rocky Moun- 
tain System, where the species is common, sepals become smaller 
and broader, petals inconspicuous or lacking, and the leaves re- 
main more or less fleshy. This austral population, the ssp. 
columbiana, extends into Wyoming and Oregon. Arenaria rossii 
seems to be absent from the Middle Rocky Mountains and the 
Wyoming Basin (both as defined by Fenneman), i.e., from the 
Uinta and Wasatch Ranges in Utah, and possibly all of Wyoming, 
except the northernmost portion. 

A taxonomic arrangement of the races of A. rossii, and a brief 
consideration of A. macrantha and its relatives, the southern 
complement of A. rossii, is provided herewith, and appended is 
a further provisional key? to the species of the Arenaria rossti— 
A. stricta complexes of America. 


ARENARIA ROSSII Robert Brown apud Richardson. 


A. rossii ssp. rossii.—4A. Rossii R. Br. apud Richards. Append. Frank- 
lin Journ. p. 738. 1823, as to Richardson Arctic specimens; A. Rossii 
R. Br., in Chloris Melvilliana p. 14. 1823, as to the Parry Voyage Plants 
of Melville Island, and “A. Rossii, Richardson in Franklin's Journ. p. 738, 
paulo diversa est statura majore, . . . "; idem, R. Br. in Suppl. Parry’s 
Ist Voyage p. 272 (celxxii), 1824; “A rossii, Brown, Supp. Parry’s Voy. 
eclxxii. No. 20. « Brown, l.c. nobis non visa.,” and “B A. Rossi Frankl. 
App. 1 ed. p. 738, No. 170," Richardson, Append, Franklin Journ. 
ed. 2 p. 745. 1823; Hooker Fl. Bor.—Am. 1: 100. 1831; Alsine rossii 
Fenzl, Verbreit. Alsin. 18. 1833; Minuartia rossii (R. Br.) Graebn. Syn. 
Mitteleurop. Fl. 51: 772. 1918; Arenaria Rossii R. Br. apud Richards. 
var. Daethiana Polunin, Bot. Can. East. Arctic Bull. Nat. Mus. Can. 
92: 201. 1940; Minuartia rolfii Nannfeldt, Nytt. Mag. Bot. 3: 161. 1954. 


In the northernmost part of its range, the subsp. ross?i is char- 
acteristically a densely and closely pulvinate plant, infrequently 
flowering, and probably often reproducing vegetatively by means 


3 For a key to the genus see: Maguire, B., Arenaria in America North of Mexico. A 
Conspectus. Am. Mid. Nat. 46: 494—498. 1951. 


46 Rhodors [Vor. 60 


of easily detached short shoots borne in the axils of the primary 
leaves (discussed at length by Nannfeldt, Le). The type speci- 
mens of Minuartia rolfii Nannf. (Simmons 2390) are of this form. 

More westerly, and perhaps generally in more protected places, 
partieularly on the mainland, the plant tends to become more 
loosely tufted and, as observed by Porsild (in correspondence) 
flowers abundantly, and presumably matures seed. The speci- 
mens of Richardson represent both forms. 

The specimens of Richardson (No. 170, the types of A. rossi) 
collected on “Barren Grounds from Point Lake to the Arctic 
Sea,” of which we have seen the material at Kew, the British 
Museum, the Gray Herbarium, the Oxford Fielding Herbarium, 
and The New York Botanical Garden, consist both of densely 
pulvinate and more loosely tufted specimens. Melville Island 
plants of the Parry Voyage, seen at the British Museum and 
Fielding Herbarium, are of the pulvinate form. 


TYPE or arctic exploration specimens seen: coast, British North 
America, Dr. Richardson 17701 1819-22, “Arenaria Rossii Br.,” (holo- 
type, BM, photo can, the holotype sheet consisting of six specimens all 
of the more compact form); second sheet with three specimens (BM); 
"Aretie America, Frankl. Exp. (GH 2 sheets, K), the tufted form; 
[Franklin Journ.] No. 170, Richardson (GH, Ny), the tufted form; 
Melville Island, *Parry's 1 Voy. No. 20. Chloris Melvilliana p. 14. 
1823." (BM, oxF), the pulvinate form. Ad Barren Valles (lat. 76° 37’, 
long. 84° 25’) sinus Harbour Fjord, Ellesmerelandiae meridionalis in 
campis argillosis, July 28, 1900, H. G. Simmons 2390 (GH, NY, isotypes 
of M. rolfii Nannf.). 

DISTRIBUTION. Barren lands at low altitude, West Greenland, the 
Arctic Archipelago and the aretie coastal mainland to the Mackenzie 
River. 


REPRESENTATIVE SPECIMENS. Greenland: Jacobsen Bay, Ymer Island, Aug. 
11. 1932, T. Sørensen 3312 (cAN); Dragon Point (82° 15’ N.), July 19, 1917, 
Th. Wulff s.n. (au). CANADA. Arctic ARCHIPELAGO. Baffin Island: Arctic 
Bay. Sept. 8-11, 1936, Polunin 2587 (aH); ? Cape Dorset, July 29, 1938. 
Manning & Manning 19 (can); Hantzsch River, Sept. 3, 1938, Manning 
& Manning 182 (cAN); Silliman Mt., July 11, 1948, Senn & Calder 3924 
(pao), with characters of ssp. elegans. Ellesmere Island: Harbour Fjord, 
July 28, 1900, Simmons 2390 (GH, NY, isotypes Minuartia Rolfii Nannf.) ; 
Craig Harbour, Sept. 16-17, 1934, Polunin 872 (can); Eggerton Lake, Aug. 
17, 1951, MacDonald 18 (can); Hawkins Lake, July 17, 1951, Bruggemann 
202 (pao); Parr Inlet, Aug. 8, 1951, Bruggemann 252 (pao); Wood Creek, 
Aug. 13, 1951, Bruggemann 262 (pao); Ward Hunt Island off n. coast Elles- 
mere I. 88° N., July 14, 1954. Christie 60 (CAN). Devon Island: Dundas 
Harbour, Sept. 7, 1936, Polunin 2554 (CAN). Prince Charles Island: Foxe 


Arenaria rossii 47 


1958] Maguire, 


Basin, Aug. 15-18, 1949, Baldwin 1939 (can). King William Island: 68° 
47’ N., 97° 40’ W., Aug. 10, 1949, Woodruff 144 (pao); Victory Point, Aug. 
8, 1954, Cooper 112 (can); Aug. 17, 1955, Cooper 151 (can). Grinnell 
Land: Lady Franklin Bay, Greely 50 (GH). Victoria Island: Wollaston 
Pen., July 27, 1949, Porsild 17216 (can); Holman Island trading post, Aug. 
8, 1949, Porsild 12278 (can); Albert Sound, Aug. 4, 1949, Porsild 17384 
(CAN). Banks Island: Cape Lambton, July 30, 1949, Porsild 17554 (can); 
n.e. corner of island, Aug. 13-20, 1949, Porsild 17667 (cAN) ; Bernard Island, 
Aug. 22, 1949, Porsild 17749 (can); De Salis Bay. July 17, 1952, Manning & 
Macpherson 16 (CAN). Cornwallis Island: Resolute Bay, July 31, 1949, 
Collins 192 (CAN) ; 75° 15’ N., 96° 20’ W., in 1952, Mackay 8 (can); Resolute 
Bay, July 30, 1949, Schofield 446 (pao). Melville Peninsula, Repulse Bay, 
July 25, 1950, Bruggemann 69 (pao). Spruce Bay, July 23, 1951, Chillcott 56 
(Dao). Kerewatin District. Southampton Island: South Bay, Aug. 22, 
1936, Polunin 2280 (au), 2282 (aH); July 9, 1948, Cody 1231B (pao); July 
25. 1948, Cody 1558 (pao); July 28, 1948, Cody 1637 (pao); July 29, 1948, 
Cody 1653 (pao); Aug. 5, 1948; Cody 1855 (pao); Aug. 11, 1948, Cody 2016 
(pao); July 15, 1948, Cody & Senn 1338 (pao); Ford River, Aug. 15, 1950, 
Brown 213 (pao). Mackenzie Districr. Bernard Harbour, Aug. 1915, 
Johansen 367 (cAN) ; Great Bear Lake, Aug. 2, 1928, Porsild & Porsild 5140 
(cAN); Tree River, July 11, 1955, Miller 94 (can); Coppermine, Aug. 4, 
1951, Findley 252 (pao). 


A. rossii subsp. elegans (Cham. & Schlecht.) Maguire, comb. nov. 
Arenaria elegans Cham. & Schlecht. Linnaea 1: 57. 1826. 

TYPE. Ad sinum St. Laurentii inter muscos et Dryadem crescens, 
Chamisso no. 13, p. 57, l.c. 

DISTRIBUTION. Coastal and montane Alaska and Yukon of the Yukon 
River drainage. Variable. Specimens with flat ascending leaves 8-10 
mm long and small flowers (sepals ca. 2.5 mm long), represented by 
A. & R. A. Nelson 4080, and L. Viereck 1400 from Mt. McKinley Na- 
tional Park, and other like specimens, probably constitute a discrete 
well segregated variety. 


REPRESENTATIVE SPECIMENS. CANADA, MACKENZIE District. Richardson 
Mts., Aug. 15-17, 1933, Porsild 6792 (can), 6793A (can); Mackenzie Range, 
Sept. 9, 1944, Porsild & Breitung 11804 (cAN) ; Canol Rd., Mile 111 east, 
July 25, 1944, Wynne-Edwards 8294 (can). Yukon Terr.: Canol Rd, 
Mile 132, June 15, 1944, Porsild & Breitung 9609 (can); Canol Rd., Mile 
105, July 21, 1944, Porsild & Breitung 10891 (CAN). Alaska. Miller House, 
115 miles n. Fairbanks, July 22-28, 1936, Scamman 168 (au); July 24, 1947, 
Scamman 482 (Gu); July 14, 1947, Scamman 4695 (an). Brooks Range, 
Jordal 3605 (can). Eagle Summit: July 7-11, 1937, Scamman 756 (an); 
July 12-28, 1940, Scamman 2074 (au); Aug. 1-9, 1940, Scamman 2247 (an); 
June 23-30, 1945, Scamman 3514 (an); July 13-15, 22-23, 1949, Scamman 
5195 (cn); June 25, 1948, Lepage 23277 (can, pao). White Mts, July 7, 
1953, Gjaerevoll 456 (can). Alaska Range, June 19, 1926, Porsild & Porsild 
225 (can). Mt. McKinley Nat. Park: Aug. 8, 1939, A. & R. A. Nelson 
4080 (cH, NY); July 17, 1956, Viereck 1400 (coro, Ny); July 10, 1956, 
Viereck 1191 (coLo, NY). Seward Peninsula: Cape Nome, in 1900, F. E. 
Blaisdell s. n. (GH, NY); Upper Kougarok River, June-July, 1909, C. B. 
Atwater s. n. (GH); Nome, Aug. 6-10, 1926, Porsild & Porsild 1336 (can, 


48 Rhodora [Vor. 60 


Gu); Nome, Anvil Hill, Aug. 6-10, 1926, Porsild & Porsild 1334 (CAN), 
1335 (CAN, GH); Bluff, Aug. 5-6, 1926, Porsild & Porsild 1222 (can); Nome, 
Anvil Hill, Aug. 9, 1948, Lepage 23895 (CAN, pao); Nome, June 20, 1951, 
Whillans 52 (pao). Norton Sound, July 16-29, 1926, Porsild & Porsild 984 
(CAN, GH). Port Clarence, Aug. 6-20, 1949, Scamman 5481 (GH). 


A. rossii subsp. columbiana (Raup) Maguire comb. nov. A. rossii 
var. columbiana Raup, Contr. Arnold Arboretum 6: 157. 1954. 

TYPE. Wet stones in sun at 5500 ft, Pass n. of Robb Lake, British 
Columbia, Mrs. J. Norman Henry 262 (GH). 

DISTRIBUTION. The petaliferous element, montane, the Stikine Moun- 
tains, Yukon, south in the Rocky Mountains to Colorado. 


REPRESENTATIVE SPECIMENS. CANADA. British Columbia. N. Kootanie 
Pass: July 29, 1883, Dawson 656A (CAN); Dawson 665 (CAN); Rainbow 
Mts., July 16, 1898, Spreadborough 19291 (can, GH); Robb Lake, July 25, 
1932, Henry 262 (aH, holotype of A. rossii var. columbiana Raup); Mt. 
Selwyn, July 19, 1932, Raup & Abbe 3951 (can, GH, NY); Raup & Abbe 
3761 (CAN, GH); Laurier Pass, July 22, 1935, Henry 718 (au); Alaska High- 
way, Mile 456, Porsild 9010 (CAN). Alberta. Moose Mt., June 29, 1897. 
Macoun 18266 (CAN, GH); July 1, 1897, Macoun 18267 (CAN) ; Crow’s Nest 
Pass, Aug. 2. 1897, Macoun 18270 (can); June 30, 1897, Macoun 268 (can); 
Saddle Mt., Banff Nat. Park, July 31, 1891, Macoun 4868 (CAN) ; Waterton 
Lake. July 11, 1931, A. S. Pease 22570 (au); Banff Nat. Park, July 13-22, 
1946, Porsild & Breitung 15886 (CAN); 15977 (CAN). 


A. rossii subsp. columbiana var. apetala Maguire, Am. Mid. Nat. 
46: 510. 1951. 

TYPE. Alpine meadow, Preston Park, alt. 7520 feet, Glacier Park, 
Montana, July 20, 1932, Maguire 732 (holotype NY). 

DISTRIBUTION. The apetalous element, forming extensive local popu- 
lations, British Columbia, Alberta, Montana, and possibly northwestern 
Wyoming. Commonly more compact, and more frequently collected 
than the preceding. A single collection is known from the Wallowa 
Mts., Oregon, and a diminutive specimen, J. T. Howell 22773 from Mono 
Mesa, Inyo Co., Calif., has seed and flower characteristics of the var. 
apetala, and is tentatively assigned here. 


REPRESENTATIVE SPECIMENS. CANADA. Yukon Terr., Canol Rd., Mile 
102, July 19, 1944, Porsild & Breitung 10615 (can). Alberta. Crow’s Nest 
Pass, Aug. 2, 1897, Macoun 18271 (cAN), 18296 (CAN) ; July 31, 1897, Macoun 
18271 (cAN). Upper Red Deer River: July 12, 1951, Porsild. 18175 (CAN) ; 
July 12, 1951, Porsild 18306 (can). Coleman, July 6, 1956, Porsild & Lid 
19351 (CAN). Mt. Inglismoldie, July 1, 1916, Lewis idi (can). Banff Nat. 
Park, Porsild & Breitung in 1945; Sulphur Mt., 12432 (can); Mt. Temple, 
12554 (CAN) ; Sunshine Ski Lodge, 13161 (CAN) ; 13423 (CAN); 13469 (CAN); 
31470 (CAN); 14102 (CAN); 15885 (CAN); Mt. Bourgeau and Mt. Brett, 
13802 (cAN); Cascade River, 14950 (can); Upper North Saskatchewan 
River, 16056 (can); Brewster Hill, 19464 (can); Citadel Mt., 19555 (can); 
Sulphur Mt., June 14, 1906, S. Brown 146 (aH, NY). Waterton Lake: 
Sheep Mt., July 28, Macoun 10098 (can); Sept. 6, 1953, Moss 10516 
(CAN); Carthew Pass, July 26, 1953, Breitung 16689 (wv). Jasper Nat. 


1958] Maguire, —Arenaria rossii 49 


Park, Medicine Lake, Aug. 7-9, 1941, Scamman 2528 (au). UNITED STATES: 
Montana. Glacier Nat. Park: Mt. Jackson, Aug. 24, 1920, Somes 70 (Ny); 
Piegan Pass, July 20, 1930, Pease 22194 (GH); Preston Park, July 20, 1932, 
Maguire 732 (xy, holotype A. rossi var. apetala Maguire); Logan Pass, 
July 16, 1934, Jones 5523 (aH). Mt. Henry, Midvale, July 16, 1903, Umbach 
405, in part (NY). Beaverhead Co.: Black Lion Mt., July 30, 1945, Hitch- 
cock & Muhlick 12908 (xy). Big Snowy Mts., July 6, 1945, Hitchcock & 
Muhlick 12037 (Ny); Pintlar Peak, July 27, 1945, Hitchcock & Muhlick 
12860 (NY). Bridger Mts., June 5, 1897, Rydberg & Bessey 4050 (ny, 2 
sheets). Big Horn Mts., July 1898, Tweedy 162 (ny); Beartooth Mts., 
Carbon Co., July 29, 1955, Cronquist 8003 (NY, GH, US, UC, WS, WTU, COLO, 
K, P). Oregon. Wallowa Mts. July 31, 1899, Cusick 2299, in part (GH). 
Wyoming. Northwestern Wyoming Expedition, in 1873, Parry 40 (aH). 


ARENARIA MACRANTHA AND ITS RELATIVES 


As shown above, Arenaria rossii, occupying the American trans- 
Arctic region and northern Rocky Mountains, apparently does 
not extend into the Middle Rocky Mountain area of Wyoming 
and Utah (the Uinta and Wasatch Ranges), or the Wyoming 
Basin. Apparently the range of A. macrantha, which replaces 
A. rossii in the south, and its relatives are confined to the southern 
Rocky Mountains and Colorado Plateau region (as defined by 
Fenneman) and do not occur in the Middle Rocky Mountain 
area. Thus, the Middle Rockies form a broad spacial hiatus 
between the ranges of the two closely related complexes. 

From the material of the complex now available, two elements 
stand out more strongly, viz., that represented by the specimens 
of the type collections of A. macrantha from Montezuma County, 
Colorado, and of A. filiorum from Iron County, Utah. 


Arenaria macrantha (Rydb.) Nels. Man. Bot. Rocky Mts., p. 186. 
1900. Alsinopsis macrantha Rydb. Bull. Torrey Club 31: 407. 1904. 

TYPE. Common alpine form, Little Kate Basin, La Plata Mts., Monte- 
zuma Co., Colorado, July 14, 1898, Baker, Earle & Tracy 678 (holo- 
type NY, isotype NY). 

The types, consisting of four plants on two sheets at The New York 
Botanical Garden, are luxuriant specimens, obviously perennial, with 
numerous procumbent stems to 10 em long; conspicuous flowers borne 
in 3 (5)-flowered cymes; sepals broadly lanceolate, 4.5-5.0 mm long, 
2 mm broad, acuminate, strongly 3-nerved; petals conspicuously ex- 
ceeding the sepals, 7-8 mm long; and leaves more or less plane, strongly 
l-nerved, blunt. No mature capsules had been formed, consequently 
seeds are lacking. Collections recently obtained show the species, as 
expected, to be somewhat variable in habit, leaf-form, and range of 
flower size. Two of them have formed mature capsules and seed. 


50 Rhodora [Vor. 60 


Adequate eireumseription is now possible: leaves may be more or less 
plane (as in the types) or triquetrous-subulate; sepals 4.0-5.5 mm long; 
petals ordinarily conspicuously surpassing the sepals; seed reniform 
ca. 1 mm broad, tesselate-tuberculate, blackish. 

DISTRIBUTION. Alpine or similar habitats, mountains of the Southern 
Colorado Rockies. 


Representative specimens, Colorado. Gray’s Peak, A. Eastwood s.n., in 
July 1888 (coro); alpine, South Park, Wolf & Rothrock 346 (au). Park 
Co., Hoosier Ridge, Weber, Rollins & Livingston 655 (coto); tundra, North 
Star Mountain, Hoosier Pass, ca. 12,300 ft. alt, Weber 8751 (coro); Bald 
Mt., 11,500 ft., Aug. 1898, E. A. Bessey s.n. (NY); Bald Mt., Aug. 28, 1954, 
Jean Langenheim 8968 (coto). Clear Creek Co. Weber & Dahl 8613, 
depauperate specimens, (coLo). Gunnison Co. Travertine bog and cliffs, 
Cement Creek Canyon, ca. 8500 ft. Weber & Langenheim 9520 (coro); 
Horse Basin, 11600 ft., Langenheim 70 (coro) ; alpine, Comanche Creek at 
12000 ft., Langenheim 1361 (coto). Pitkin Co. ridge east of Avalanche 
Creek, 12000 ft., Langenheim 2106 (coto. NY). Montezuma Co. Little 
Kate Basin, Baker, Earle & Tracy 678 (holotype Ny, isotype Ny). San 
Juan Co. Near Irontown, July 21-31, 1899, C. C. Curtis sn. (NY). 


Hall & Harbour No. 69, Lat. 39?—41?, Colorado, in 1862 (GH), 
an apetalous form with 1—3-flowered eymes and reddish slightly 
sculptured seed, can hardly be assigned to A. macrantha. It is 
similar to a few scattered collections obtained from the Northern 
Rocky Mountains and Canadian Rocky Mountains (viz.: Porsild 
& Breitung 10615 (CAN), an apetalous form from Mile 102, Canol 
Road, Yukon Terr.; and Spreadborough 19290 (can), Rainbow 
Mt., Fraser River, B. C., petalous form), which are rare, sporadic 
and do not form populations, and have for the time being been 
assigned to A. rossii subsp. columbiana. 


Arenaria filiorum Maguire, Bull. Torrey Club 73: 326. 1946. 

TYPE. Common, gravelly beach, Navajo Lake, Iron County, Utah, 
July 13, 1940, Maguire 19472 (holotype Ny, isotype GH, UTC, UC). 

Small glabrous annual from a slender taproot, with cymes usually 
bearing 3-5 flowers; sepals are ovate-lanceolate, strongly 3-nerved, and 
are 3.5-4.8 mm long; petals more or less equaling or shorter than the 
sepals; seed abundantly produced 0.7-1.0 mm broad, reniform, very 
dark reddish brown or blackish, rather strongly sculptured (in con- 
trast to the seed of A. rossii). 

This highly distinetive form occurs intimately with A. rubella, where 
both hold dominance on the gravelly beach of the lake. Quite similar 
plants, L. Ellison 4523 (ny), Island Lake, San Pete County, were col- 
lected also on gravelly lake beaches. 

Elsewhere from the high Colorado Plateau of Utah, viz., Maguire 
19988 (NY), 10928 ft. alt., Mayfield Canyon, and Maguire 20060 (NY), 
12000 ft. alt., Horseshoe Mt., both in San Pete County, and Maguire 


Arenaria rossii 51 


1958] Maguire, 


20097 (xy), East Brian Head Peak, 11000 ft. alt., Iron County, are 
somewhat similar plants but obviously perennials. They most closely 
resemble small-flowered members of the complex cited above from 
Colorado, but are distinguishable in minor ways from them. 

A series of specimens obtained from the Charleston Mts., Clark Co., 
Nevada, Clokey 5460 (NY), 7510 (NY), and 7923 (NY), collected in 
1935, 1937 and 1938 respectively, are obviously perennial with commonly 
uniflorous cymes. They were initially assigned to A. filiorum but are 
easily recognizable as distinct from it. They are extremely uniform 
and certainly form a geographically restricted race, yet undoubtedly 
belong to the A. macrantha-filiorum complex. 


A KEY‘ TO THE ARENARIA ROSSII-À. STRICTA? COMPLEXES AND 
THEIR RELATIVES IN NORTH AMERICA 


1. Seed reniform (with respect to the hilum), hence broader 
than long; plants completely glabrous. 
2. Seed smooth or inconspicuously ornamented (under X 10 
magnification), pale, reddish, 0.5-0.7 mm broad, cymes 
üniflorous. ebracteate. L. e yen see RE cesse ele A. rossit. 
3. Sepals commonly 1.5-2.5 (3.0) mm long, oblong-ovate, 
obtusish, usually 1-nerved ;° petals conspicuously sur- 
passing the sepals; cymes uniflorous, ebracteate ; 
pedicels 5-20 mm long; leaves fleshy, subulate, obtuse, 
2-4 mm long; densely pulvinate or tufted plants of 
(Spitzbergen and) Arctic America; Greenland, Arctic 
Archipelago, and Arctic Coastal America to the 
Mackenzie River Region. ............... A. rossii subsp. rossii. 
3. Sepals commonly 3.0-3.5 mm long, lanceolate, acute, 
commonly 3-nerved; pedicels 2-4 (5) em long; leaves 
less fleshy, more or less plane, acutish, petals nar- 
rower, equaling or commonly somewhat shorter than 
the sepals; loosely tufted plants of Alaska, Yukon in 
the Yukon River drainage, and probably the Richard- 
son Mountains. ......... ey TO A rossii subsp. elegans. 


4 This “key” is intended to supplement, by introducing some corrections and additions, 
the key offered by me in 1951 (1.c.). In no sense do I mean to imply that the species 
herein admitted are necessarily more strongly intra-related, than to or with other members 
of the Sect. Alsine. 

^ Dr. A. E. Porsild in correspondence of Aug. 6, 1957, wrote, “I wonder if you are not 
overlooking that Minuartia stricta (Sw.) Hiern. is not the same as Arenaria uliginosa 
Schleich and certainly not synonomous with A. stricta ssp. dawsonensis as suggested by 
vou (1951)." Both from lack of material and opportunity to extend the present study, 
I cannot have a competent self-gained opinion on the point. In view of Dr. Porsild’s 
broad knowledge of boreal American plants, and his careful study, I am quite content 
to accept his interpretation of plants so indicated by him. Obviously, this would require 
the addition of A. uliginosa to my “key” of 1951; and indeed also A. macrantha (Rydb.) 
Nels., there inadvertently omitted by me. 

8 Anatomically the sepals in the entire complex are 3-nerved. The lateral nerves may 
be prominent, or weakly developed. Or, the lateral nerves may be obscured, in more 
crassulous sepals, by relatively thicker mesophyll tissues, thus in appearance the sepal 
becoming ''1l-nerved." The terms ''3-nerved" or ‘‘l-nerved’’ merely give expression for 
relative prominence of the lateral nerves. 


52 Rhodora [Vor. 60 


3. Sepals 2.5-3.5 mm long, lanceolate or ovate-lanceolate, 
acutish, moderately or weakly 3-nerved, or often 1- 
nerved; petals inconspicuous, narrow, shorter. than 
the se pals, often lacking; leaves more or less fleshy, 
triquetrous or subulate; Rocky Mountains from the 
Stikine Mountains south to Oregon and northern 


Wyoming. ....... ne beeacdcsereares A. rossii subsp. columbiana. 
4. Petals present ........00. 0000. eos Var. columbiana. 
4. Petals lacking ........... Lee cece cnet eee e eee . var. apetala. 


2. Seed obviously tesselate- tubercul: ate (under X 10 magni- 
fication), blackish, (0.8) 1.0-1.5 mm broad; cymes com- 
monly 3-several-flowered, bracteate; sepals strongly 3- 
nerved. 
5. Stems mostly under 10 cm high (or long), the 
inflorescence shorter than the vegetative portion 
of the stem, cymes commonly 3-5-flowered. 
6. Plants glabrous. 
7. Obviously perennial; cymes 1-3-flowered, 
flowers large, showy; sepals 4.0-5.5 mm 
long, lanceolate, acute; petals commonly 
exceeding the sepals, to 8 mm long; plants 
rather densely tufted; mountains of cen- 
tral and southern Colorado and the type 
collection from the La Plata Mountains, 
Colorado. ....... Sv sseeveesecees Arenaria macrantha. 
7. Annual or weakly perennial; flowers not 
showy; cymes 3-7-flowered; sepals 4.0-5.5 
mm long; petals shorter than the sepals; 
apparently restricted to the high Colorado 
Plateau of southcentral Utah. .......... A. filiorum. 
6. Plants conspicuously glandular; circumboreal. 
en eee ess A. rubella. 
5. Stems 10- 30 cm ‘high: primary leaves 1-2 cm long; 
inflorescence characteristically exceeding the 
hyaline portion of the stem; sepals 3.5—5.0 (6.5) 
mm long, 3-nerved, rarely 1-nerved; petals 5-8 
mm long; seed 0.8-1.5 mm broad. .............. A. stricta. 
8. Plants totally glabrous, strongly perenni: il. 
9. Petals conspicuously surpassing the 
‘talyx; capsule equal to or shorter 
than the calyx. 
10. Stems leafy for more than half their 
length; primary leaves usually 
1.5-3.0 em long; plants mostly 
lax, frequently matted. lv 
VENE es eeebrerivie A. stricta subsp. stricta. 
10. Stems leafy usually below the mid- 
dle; primary leaves mostly 0.5-1.5 
cm long; plants rigid, not matted. 
ode eseesvarsoetsouves A. stricta subsp. texana. 
9. Petals equal to or mostly shorter than 


1958] Maguire,—Arenaria rossii 53 


the calyx; capsule exserted. ........ 
AU EUER COE A. stricta subsp. dawsonensis. 
8. Plants glandular-pubescent, at least in the 
inflorescence, weakly perennial. ....... 
DPA RNC A. stricta subsp. macra. 
1. Seed oblong (with respect to the hilum), hence longer than 
broad. 
11. Primary leaves 3-nerved, non-glaucous; inflorescence 
stipitate-glandular, half or less than half the length 
of the glandular stem ; sepals 3.5-6.5 mm long; ovules 
9-15; seed several or solitary, oblong, 1.0-1.3 mm 
broad, 1.25-20 mm long; widespread in the Rocky 
Mountain, Intermontane, and Cascade-Sierra Nevada 
region of western America, in the mountains from 
British Columbia and Alberta to California, Nevada, 
Utan and Calorado a6.) eg ae DEM ee A. nuttallit. 
12, Leaves ascending or strict, neither arcuate nor squar- 
rose. 
13. Leaves abruptly acute or apiculate, infrequently 
pungent; sepals acuminate, midrib not promi- 
nent. 
14. Petals shorter than the sepals; Cascade 
Range, Rocky Mountains. .. A. nuttallii subsp. nuttallii. 
14. Petals longer than the sepals; north Cali- 
fornia coast ranges, Siskiyou Mountains. .. 
A Pees me ra eer nen yee A. nuttallii subsp. gregaria. 
13. Leaves strongly pungent; sepals narrow, pun- 
gently attenuate, midrib prominent; Sierra 
Nevada Range. ............... A. nuttallii subsp. gracilis. 
12. Leaves arcuate or squarrose. 
15. Sepals 3.5-5.5 (6.0) mm long, lanceolate, 
sometimes broadly so, acuminate, 1- 
nerved (occasionally 3-nerved); petals 
shorter than the calyx; Cascade Range, 
Rocky Mountains. ...... A. nuttallii subsp. nuttallii. 
15. Sepals 5.5-6.6 mm long, narrowly lanceo- 
late, pungently attenuate, 3-nerved; 
petals more or less equaling the calyx; 
central Great Basin. .... A. nuttallit subsp. fragilis. 
11. Primary leaves 1-nerved; glaucous; inflorescence diffuse, 
exceeding the length of the glabrous stem; sepals 3.0— 
4.5 mm long; ovules 6 (7); seed solitary, 1.4-1.5 mm 
broad, 2.3-2.8 mm long; plants of serpentine areas, 
Coast, Ranges, Trinity County, California. ............. A. roset. 


54 Rhodora [ Vor. 60 


RUMEX STENOPHYLLUS IN NORTH AMwEniCA.— l'wo. specimens 
of Rumex stenophyllus Ledeb. have been discovered in a collection 
of plants from Manitoba. They were found growing in a road- 
side diteh not far from the village of Otterburne, about 30 miles 
SE of Winnipeg, and were collected by the junior author on 
Aug. 21, 1950. 

Rumex stenophyllus Ledeb. is a continental Eurasiatic species, 
native to eastern and western Siberia, central Asia, eastern 
and middle Europe to lower Austria, Moravia and central 
Germany (Rechinger 1949). It is occasionally introduced 
into Scandinavia (ef. Hylander 1955), Holland (ef. Rechinger 
1949), and England (cf. Clapham, Tutin & Warburg 1952). The 
most favorable habitats are roadside ditches and other places, 
where the soil is slightly saline and occasionally flooded. At 
times the plant spreads to road shoulders and wastelands, 
or may even become a field weed. 

'The Red River valley, where the plant was found, was origin- 
ally settled by French-Canadian farmers, but early this century 
a great influx of immigrants came from the Ukrainian parts 
of central and eastern Europe. Like other immigrants, these 
Ukrainian farmers brought their own grain, which included 
an assortment of weeds. Although this alien flora still is not 
fully known, it includes some very well naturalized species, 
like e.g. Sonchus uliginosus M. B., which is now widespread 
all over the marshes of central Canada and the adjacent United 
States. Rumex stenophyllus apparently also belongs to this 
group of species, although its occurrence is not as evident as 
that of the conspicuous Sonchus. Only a single locality of 
Rumex stenophyllus is known so far from Manitoba, but the plant 
is undoubtedly more widespread. The valleys on the prairies 
with their slightly saline and repeatedly flooded soils certainly 
offer conditions very much like those of its original habitat. 

Aceording to Frankton (1955, and in litt.) the species is not 
uncommon in Saskatchewan, where it was first discovered a few 
miles north of Swift Current in 1954, when seed collections were 
sent in for identification by Mr. A. Budd. In the herbarium 
of the Dept. of Agriculture, Ottawa, specimens from the fol- 
lowing states have also been uncovered: Colorado, Nebraska, 


Wyoming, N. Dakota, S. Dakota and Minnesota, under the fol- 


1958] Lóve and Bernard,—Rumex stenophyllus 55 


lowing names: R. obtusifolius, brittanica, pulcher, alluvius and 
odontocarpus. R. odontocarpus (cf. Moore, 1957) and R. alluvius 
(cf. Gates & McGregor, 1950; Rechinger, 1952) are synonyms 
to R. stenophyllus. 

Due to the fact, that the species has been the object of so much 
confusion, it seems appropriate to repeat in translation the de- 
tailed description given in latin by Rechinger (1949): 


"Root fusiform, vertical, perennial (occasionally the plant flowers during 
its first year?), stem stiffly erect, 20-60 (-120) em. tall, brownish or reddish, 
suleate-striate, leafy, often divided above the middle into several branches, 
but in depauperate specimens sometimes subsimple. Branches erect, or 
erect-divergent, very rarely arching; in sturdy specimens lower branches 
sometimes + fasciculate and repeatedly branched, but usually all branches 
straight and simple, forming an open, but narrow panicle. Basal leaves 
lanceolate, acute, their bases broadly or narrowly cuneate, their tips gradually 
attenuate, their laminas flat or with slightly undulate margins, the leaf stalks 
+ equal to or rarely longer than the blades. Stem leaves lanceolate, nar- 
rowing towards both ends, most of them with flat margins but some slightly 
crenulate-crispate towards the tip, 4-6 (-7) times longer than broad, smooth 
and glabrous, with the lateral nerves leaving the midnerve at 40-60? angle. 
Upper leaves with increasingly shorter leaf stalks and topmost leaves + linear. 
Flowers in many-flowered whorls, the lower ones + remotely leafy, the upper- 
most almost continuous and perfectly aphyllous. Fruit pedicel narrowly 
filiform, of varying length, but often 1.5-2 times as long as the mature perigone, 
and in its lower 3rd or 4th part with a distinct ring-joint. Close to the perigone 
the pedicel widens to funnelshape, or becomes at least considerably thicker. 
Outer perigone leaves linear-lanceolate of about half the length or slightly 
shorter than the valves, to which their margins are pressed. "Valves cordate 
triangular, the base slightly cordate or subtruncate, the tip acute, membra- 
naceous, the surface subregularly reticulate with slightly raised veins, the 
areas between the nerves elongate towards the margins, which form more or 
less irregular, sharp and coarse teeth, +0.5, rarely 1, mm. long, on both sides 
of the entire-margined tip. Valves (3.5-) 4 (-rarely 5) mm. long, about as 
broad or rarely somewhat narrower, all with about equally large grains. 
Grain golden-brown, almost smooth, ovate-ellipsoid, very prominent, the 
aeute tip often running out into and continuing in the median nerve of the 
valve. The grain is about half the length of the valve and occupies 4 to 4 
of its width. Mature nut dark brown, about 2 mm. long and 1.5 mm. broad, 
apiculate at both ends, the tip somewhat sharper than the base." 


Superficially, Rumex stenophyllus resembles R. crispus, but 
there are several significant differences. Rechinger (1949) 
points out, that R. stenophyllus possesses two characteristics 
which never occur in true R. crispus, namely valves with dis- 
tinctly toothed margins and a valve-nervature which forms 
elongate, flat meshes (cf. fig. 1). The two species were placed 
in very different sections and sub-sections by Rechinger (1949) 
and Losina-Losinskaja (1935). 


56 Rhodora [Vor. 60 


Although Rumea stenophyllus was described as a species from 
Altai by Ledebour in 1830, its occurrence in Europe was obscured 
for a long time by its casual resemblance to R. crispus and 
R. obtusifolius. It has thus been regarded as a variety of these 
two species, i.e. R. crispus var. dentatus Schur. or R. obtusifolius 
var. cristatus Neilr., or even as only a hybrid between them, 


Fia. 1: a, Rumer stenophyllus Ledeb., b. Rumex crispus L, 


k. crispus X obtusifolius. Other authors have regarded it 
as a separate species, R. biformis (Menyh.) Borbas, or R. odon- 
tocarpus Sandór (cf. Mansfeld 1940), thought to be endemic 
in central and eastern Europe. However, in all recent European 
manuals, the plant is identified with the Altaian taxon, since 
the type material of this species does not differ from European 
material (cf. Rechinger 1949). 

Rumez stenophyllus differs from the real hybrid R. crispus X 


1958] Lóve and Bernard,—Rumex stenophyllus 57 


obtusifolius in several morphological characteristics, e.g. in the 
shape of leaves and valves. However, the hybrid is always 
sterile, and the species is fully fertile. "The sterility is caused 
by the fact, that the hybrid has 2n — 50 chromosomes, its 
parents 2n — 60 and 40 chromosomes, respectively (Lóve 1942). 
R. stenophyllus, however, has 2n — 60 chromosomes, as de- 
termined by Pólya (1950) on Hungarian material and confirmed 
on germinating seeds of the Manitoba plants by Mrs. N. Sarkar 
(unpubl.). There is no reason to suspect, that any of the 
R. crispus X obtusifolius hybrids reported from North American 
localities by Rechinger (1937) belong to R. stenophyllus. 

A specimen of Rumex stenophyllus Ledeb. has been deposited 
in the herbarium of the Institut Botanique, Université de 
Montréal.—Doris LóvE AND FRÈRE JEAN-PAUL BERNARD, 
O.S.V., INSTITUT BOTANIQUE, UNIVERSITÉ DE MONTRÉAL AND 
INSTITUT DES SOURDS-MUETS, MONTRÉAL. 


LITERATURE CITED 


CLAPHAM, A. R., Turin, T. G. & Warsure, E. F. (1952) Flora of the 
British Isles —Cambridge. 

Frankton, C. (1955) Research Report, Canada National Weed Com- 
mittee, Western Section, pp. 123-124. 

Gates, F. C. « McGrecor, R. L. (1950) Rumex alluvius, a new 
species of dock from Kansas.—Trans. Kans. Acad. Sci. 53: 186—189. 

HLANDER, N. (1955) Förteckning över Nordens växter I. Kärl- 
vüxter.—Lund. 

LEDEBOUR, C. F. (1830) Flora Altaica 2.— Berolini. 

Love, 'A. (1942) Cytogenetic studies in Rumex III. Some notes on 
the Scandinavian species of the genus.—Hereditas 28: 289-296. 
LosiNA-LosINSKAJA, A. S. (1935) Rumex in Flora SSSR V: 444-482. 
MANSFELD, R. (1940) Verzeichnis der Farn- und Bliitenpflanzen des 

Deutschen Reiches.—Ber. Dtsch. Bot. Ges. 58 a: 1-323. 
Moorr, J. W. (1957) Notes on flowering plants in Minnesota.— 
Rwopora 59: 6-8. 
Pórnva, L. (1950) Magyarországi növényfajok kromoszómvaszámai 
II.—Ann. Biol. Univ. Debr. I: 46-56. 
RecuinGer, K. H. (1937) The North American species of Rwmexr.— 
Field Mus. Nat. Hist. Vol. XVII, 1: 1-151. 
(1949)  Rumices asiatici. Vorarbeiten zu einer Monographie 
der Gattung Rumer. VII.—Candollea 12, July 1949: 1-152. 
(1952) Beitrüge zur Kenntnis von Rumex. X.—Osterreich. 
Bot. Zeitschr. 99, 4: 523-527. 


58 Rhodora [Vor. 60 


ELATINE TRIANDRA IN NEW Yonk.—Zlatine triandra Schkuhr 
previously was found in the Manual Range in several localities 
in Wisconsin and in Skowhegan, Maine, (Fasset, RHODORA 41: 
370. 1939). I can now report the species (forma submersa 
Seubert) growing spontaneously in a swamp in Brooklyn, New 
York City (voucher deposited in The New York Botanical 
Garden, Monachino 612, Prospect Park, near the Music Grove, 
several dense mats in soft mudbank and shallow water of small 
over-grown swamp, Aug. 29, 1957; all submerged after rain, 
Sept. 14). It was confined to several patches in one small area, 
but thriving vigorously. The stems measured were up to 30 em. 
long, the internodes 1 em.; the opposite scarious-stipulate leaves 
oblong-lanceolate, up to about 1 em. long, 3 mm. wide, lightly 
emarginate at apex; the inconspicuous flowers trimerous; the 
green tiny depressed-globular fruits plentiful though solitary in 
the leaf-axils, sessile, 1 mm. high, 1.4 mm. broad, the placenta 
extending half way up the capsule; the minute, numerous, slightly 
arcuate seeds (not fully matured) radiating from the central pla- 
centa, the upper ones erect, the lower spreading, about 0.5 mm. 
long, their angular pits about 24 in each row. 

The swamp is fed by a brooklet and flows into Prospect Park 
Lake a short distance away. The yellow floating-heart, Nymph- 
oides peltatum, and Sagittaria latifolia were the most conspicuous 
plants near the Elatine. Other aquatics nearby were Chara, 
Hydrodictyon, Potamogeton, Lemna. Further off were seen a 
few plants of Mimulus guttatus. The center of the swamp was 
choked with rice cut-grass. There were large tangles of Solanum 
dulcamara. With Bidens frondosa was growing the form of 
B. connata with leaves mostly simple to tripartite. Several other 
plants but sparsely represented were Erechtites, Lythrum, Rumex, 
Polygonum, Glyceria, Echinochloa, Panicum. The shrubs in the 
swamp were Cephalanthus, Sambucus, Cornus, Salix. It is 
reasonable to suppose that the Elatine is adventive, possibly 
introduced with the European floating-heart. Quite suggestive 
of this is the fact that the species (examined, but no complete 
specimen collected) also was growing in the lily pool in the 
Brooklyn Botanie Garden. 

The discovery of a second eastern station for the waterwort 
(previously collected more than a quarter of a century ago in 


1958] Rollins, —Species Plantarum 59 


only one place in Maine) is not too surprising. "There is a likeli- 
hood that this unobtrusive weed is more extensively distributed 
than records show. However, how frequent it is in our ponds and 
other fresh water sites remains to be demonstrated. The water- 
wort has much the aspect of the water starwort, until the details 
are examined closely. The habit, shade of green, and the leaves 
even as to their retuse tips are similar in the two aquatics. Of 
course there are vegetative differences: in Callitriche the leaves 
at the end of the branches are often crowded with the upper re- 
duced to give a rosette or star-like appearance, they have char- 
acteristic punctation and are not stipulate; in Elatine the leaf- 
margins are obscurely and remotely punctiform-crenulate. Al- 
though it is generally essential in latine to study the seeds to 
determine the species (or varlety, as understood by Fasset), the 
leaf-size of our plant is not described for any other species or 
form in America but E. triandra f. submersa.—Jose?H Mowa- 
CHINO, THE NEW YORK BOTANICAL GARDEN, N. Y. 


SPECIES PLANTARUM—MoRE THAN a FacsrMILE EprrroN,—This is 
a “curious” book—if we interpret “curious” to mean “interesting,” 
following the usage of Gronovius in his reference to the famous Hortus 
Cliffortianus, interesting and also important to the systematic botanist, 
who, even today, must constantly refer to the works of Linnaeus. 
William T. Stearn of the British Museum has done an excellent job of 
bringing together in an extensive introduction the pertinent material 
to make not only the works of Linnaeus fully understandable but also 
the nature, use and limitations of the Linnaean herbarium in typification 
procedures. Furthermore, he has made clear the bearing and inter- 
dependence of contemporary books and other works of Linnaeus on the 
Species Plantarum. The introduction to Volume I is a treasure of vital 
information for the student and scholar alike. Perhaps some conception 
of the range of coverage may be seen from a mere listing of the chapter 
headings. These are as follows: The nomenclatural importance of the 
Species Plantarum; Principal events in the life of Linnaeus; Major 
botanical publications of Linnaeus; Linnaeus’s sexual system of classi- 
fication; The Genera Plantarum and the tvpification of Linnaean genera; 
Hortus Cliffortianus; The Amoenitates academicae and the authorship 
of Linnaean Dissertations; The preparation of the Species Plantarum 

! Species Plantarum by Carl Linnaeus. A Facsimile of the first edition, 1753. Volume 
I, with an introduction of 176 pages by W. T. Stearn. Publication No. 140 of The Ray 
Society, London, 1957. Sold by Bernard Quaritch Ltd., 11 Grafton St., London, W. 1, 
£2, S 10. 


60 Rhodora [Vor. 60 


and the introduction of binomial nomenclature; The reception of the 
Species Plantarum in England and its influence on British botany; 
Sources, format, method and language of the Species Plantarum; Edi- 
tions and variants of the Species Plantarum; Geographical names in 
the Species Plantarum; The species-concept of Linnaeus; Signs used by 
Linnaeus; General Bibliography. A facsimile of the 560 pages of Volume 
I of the Species Plantarum plus an added index to genera and classes 
completes the volume. The value of the index is increased by the in- 
clusion of page references to the 5th edition of Genera Plantarum, with 
which the names of Species Plantarum are to be associated. 

Many points concerning the herbarium materials used by Linnaeus 
are dealt with in a clear and concise way. Heretofore, much of this 
information could be learned by the student or young scientist most 
effectively from older persons through a kind of apprenticeship asso- 
ciation with them. Although much information had been written down 
in scattered places, a great deal of it was essentially unavailable. Part 
of it got passed on from one generation of botanists to another by 
word of mouth. This situation resulted, in part at least, from the very 
complex history of the Linnaean herbarium itself, as well as the other 
collections that figured in the typifiecation of species described by 
Linnaeus. This history has only gradually been pieced together. Now, 
this situation is well taken care of by the material at hand in the very 
fine volume under review. Assiduous study of the introductory material 
in this volume should be a must on the list of every young botanist, 
not to mention some of us who are not as voung. Fortunately, the price 
of the volume is relatively modest and the workmanship on the book 
itself is of good quality. Many botanists will want to own personal 
copies I am sure.—R. C. RorriNs. 


Volume 60, number 709, including pages 1—32, was issued 27 February, 
1958. 


P 
X47 


Lae" 


Hovova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS l 
RALPH CARLETON BEAN 
CARROLL EMORY WOOD, JR. 


( Associate Editors 
IVAN MACKENZIE LAMB 


Vol. 60 March, 1958 No. 711 
CONTENTS: 
The Spontaneous Flora of an Old House-Lot in Eastern 
Massachusetts. Richard J. Eaton ............. cee ee ees 61 
Thelepogon elegans Roth in the Dominican Republic. 
PES UU lg ko aes acs ¥ ee 04,0056 4 cee kas ce ee 73 
The Fruticose and Foliose Lichens of Worcester County, 
Massachusetts. Vernon Ahmadjian...........00c cece eens 74 


Three Changed Authorities for Combinations. Robert C. Foster.. 76 
Some Identities in Halesia (Styracaceae). R. K. Godfrey........ 86 


Che New England Botanical Club, Jne. 


8 and 10 West King St., Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


RHODORA.—A monthly journal of botany, devoted primarily to the 
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CARD-INDEX OF NEW GENERA, SPECIES AND 
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GRAY HERBARIUM of Harvard University, 
Cambridge 38, Mass., U. S. A. 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 March, 1958 No. 711 


THE SPONTANEOUS FLORA OF AN OLD HOUSE-LOT IN 
EASTERN MASSACHUSETTS 


RICHARD J. EATON 


For several years, and intensively during 1955-56, I have at- 
tempted to collect specimens of all the vascular plants to be found 
growing spontaneously on the three-acre plot of land surrounding 
our early eighteenth century farm house in Lincoln, Massachu- 
setts. The census is reasonably complete as of November, 1957, 
although subsequent additions are to be expected either as species 
previously overlooked or as new arrivals. In this latter category 
several recent. instances have come to notice, thus giving striking 
evidence of the dynamic status of local floras. For example, a 
seedling of the rapidly spreading Rhamnus Frangula appeared in 
my strawberry bed early in 1954 where evidently it had been 
“planted” by a bird. Aster novae-angliae arrived in my “meadow” 
as two inconspicuous seedlings in 1952. A third instance is par- 
ticularly interesting. A vigorous fruiting specimen of Vitis aesti- 
valis emerged from a dense shrubbery behind my tennis court in 
1956, festooning the wire backstop. I am convinced that there was 
no grapevine in the shrubbery five years earlier when I last gave it 
a thorough pruning. The nearest known station for this locally 
rare grape (where it is unquestionably indigenous) is about three- 
quarters of a mile to the northeast on a trap outcropping in asso- 
ciation with the locally rare Aster infirmus. 

The analysis of the spontaneous vegetation of this particular 
piece of land may be of scientific interest for several reasons. The 
present house with attached small barn was built about 1720. 


62 Rhodora [Vor. 60 


Presumably, the house lot was cleared before that time, possibly 
much earlier. Judging from the apparent age of a few surviving 
ancient apple trees and from pictures taken about 1880, the easterly 
half of the lot probably has been an unplowed orchard for at least 
one hundred years. Its surface soil is relatively poor and un- 
attractive for tillage. Vertical sections show vague color stratifi- 
‘ations above the depth reached by a plow, a condition which 
strengthens my guess that this portion of the lot never has been 
plowed. It may have been pastured in the early years, and when 
not pastured surely mowed annually for hay. There is hearsay 
evidence that the “meadow” (15 acre +, described below) was 
used as a truck garden for an indeterminate number of years prior 
to 1925. Thus, ecologically speaking, one-half of the lot may have 
been partially shaded sod land continuously from the time it was 
originally cleared more than two hundred years ago, its superficial 
soils varying from thin sandy loam to moist deep clayey loam. 

By making a reasonably convincing estimate of the composition 
of its original climax forest and by analyzing the present vegeta- 
tion, it should be possible to make an educated guess as to what 
species in that forest may have survived in situ down to the present 
day. We can also indicate what native species may have invaded 
the area after it was cleared and still persist under existing ecolog- 
ical conditions. Of equal interest, perhaps, is the role played by 
the introduced and adventive old world species in competition with 
our native flora in a “civilized” habitat which has remained more or 
less stable for two centuries or more. 

Other questions of interest emerge: What forest species in- 
digenous to Lincoln tend to “seed in" regularly only to be de- 
stroyed by scythe or fire? (Controlled burning of the orchard 
grassland in December or early spring has been my usual practice 
since 1940.) What native species appear only in the artificial 
habitat of leaf mold and mulch under the shrubberies and border 
plantations? What weedy species conspicuously resist the severe 
conditions imposed on them, and what ones are conspicuous by 
their scarcity or absence? In this latter connection, T find that 
Solidago ssp. (notably canadensis and rugosa which were dominant 
in the “meadow” in 1940) survived two mowings annually but 
have nearly disappeared when a three-mowing schedule was in- 
augurated. 


1958] Eaton,—Spontaneous Flora in Eastern Mass. 63 


DESCRIPTION OF THE LOT 


The three-acre lot, more accurately 2.75 acres, is a squarish 
trapezoid bounded on three sides by stone walls and on the east by 
the embankment of the town pressure reservoir. It is situated 
just under the crest of the highest land in Lincoln with a west- 
northwest exposure. High deciduous shade trees, young hemlocks, 
and shrubberies line the south and north boundary walls, with 
numerous trees of varying ages elsewhere, including a very large 
vigorous white pine adjacent to the barn on the north side. A 
very small area in the northeast corner has been allowed to grow 
up undisturbed for at least thirty years, as judged by the appear- 
ance of several hickory trees along the wall. The lawns, house 
site, garden, and tennis court occupy, roughly, the westerly half 
of the lot along the street. The easterly half is still primarily an 
orchard of mostly young fruit trees. From among the ten ancient 
apple trees standing in 1940 two survive. Thus, portions of the 
orchard are open sunny hillside. Its turf is dense and in many 
places interlaced with a close mesh of poison ivy runners. 

The hill, except possibly its core, is of glacial origin, probably a 
modified drumlin, with the usual erratic boulders and underlying 
till. A moderate northwesterly slope dips into a shallow draw, with 
a gentle rise to the street. The drainage is northerly along this 
draw and includes effluent from a septic tank. Formerly, I am 
reliably informed, this draw was a narrow swamp extending to a 
tiny pond hole south of the property. It was drained many years 
ago by a covered ditch. Today part of this swampy area is the 
site of a “meadow”. I have made a small plantation of spruce, 
hemlock, larch and native deciduous trees in moist rich soil on the 
northerly side of the meadow, thus providing a suitable habitat 
for several locally adventive species not present in 1940. Additions 
may be expected in future years when the aggressive species of 
sunny grassland completely fade out. Adjacent to it is a vegetable 
garden. The soil there is a deep clayey loam, usually moist even 
during very dry summers. Elsewhere on the hillside the surface 
soils vary from their somewhat sterile sandy loam to moderately 
deep clayey loam. In general, except where locally modified by 
applications of ground limestone, they are acid, varying slightly 
from pH 4.95 in the draw to pH 4.90 at the top of the orchard. 


64 Rhodora [Vor. 60 


CULTURAL MANAGEMENT IN RECENT YEARS 

Since 1940, the hillside orchard has been mowed once annually 
in June or July, and the meadow at least twice until 1950. Since 
then the lush growth of the latter plot has required three mowings 
each year. Excluding the war years, each area has been burned, 
generally in December or early spring, except under the fruit trees 
and narrow strips along the property lines. This latter practice 
tends to check ubiquitous poison ivy, and numerous herbaceous 
weeds, such as wild carrot, goldenrod, fleabane, yellow daisy, and 
dandelion. By burning on calm sunny days when the soil is cold 
and moist the slow fire seldom if ever reaches down to the erowns of 
the herbaceous plants and apparently has no adverse effect on seeds 
in the very top layer of the soil. Parenthetically, no burning was 
done in the springs of 1956 or 1957, nor was the orchard mowed 
until after the maturity of most of the grasses later in the season, 
in order to make collections in suitable condition for accurate de- 
termination. Weeding of the shrubberies and elsewhere was de- 
ferred for the same reason. "The young fruit trees are heavily 
mulched with hay, and the shrubberies with leaves and compost. 
The garden is fertilized with spent compost from a mushroom farm 
and sparingly with chemicals, including ground limestone. It is 
cultivated regularly after each rain, thus accounting for the paucity 
of the usual garden weeds and for the complete absence of a few 
common species. 

The vegetation in the meadow is now subject to considerable 
artificial control to improve the hay crop. Fifteen years ago the 
grasses were in severe competition chiefly with a large and rapidly 
expanding patch of Helianthus tuberosa, with Solidago rugosa and 
S. canadensis, Asclepias syriaca, [quisetum arvense, and other 
"weeds" in the descending order of abundance. Today, the 
chiefly a few lush species from the Old. World—are 


grasses 
dominant. 
ANNOTATED CHECK LIST 

The following lists of plants found on the house lot are based on 
collected. specimens deposited in the Herbarium of the New 
England Botanical Club. I have excluded all species which have 
been deliberately introduced by myself or presumably so by my 
predecessors, including spontaneous seedlings from introduced 
shade trees on the place or in the neighborhood, such as Juglans 


1958] Eaton,—Spontaneous Flora in Eastern Mass. 65 


nigra, Betula alba, Acer platanoides, A. saccharum, A. saccharinum, 
and Catalpa bignonioides (of which a single seedling appeared for 
the first time in 1956). The nomenclature and sequence of families 
and genera follow Gray's Manual, 8th Ed. Species within a genus 
are listed alphabetically for convenience, authors’ names being 
omitted for the sake of brevity. 


TABLE 1 


SPECIES INTRODUCED AND NATURALIZED FROM THE OLD WORLD 


Bromus commutatus. Scarce. First observed in 1956. Vineyard. 
Festuca elatior. Common. Variable. Meadow and orchard. 

F. ovina. Scarce. Rocky, sterile soil on retaining wall. 

Poa compressa. Scarce. Dry, stony soil, edge orchard. 

Dactylis glomerata. Common. Meadow and orchard. 

Lolium perenne. Lush turf. Near edge of lawn. 

Arrhenatherum elatius f. biaristatum. Scarce. Vineyard. 

Agrostis alba. Abundant in moist ground. 

A. canina f. mutica. Several large colonies. Sunny orchard. 

A. tenuis. Rather scarce. Moist sunny orchard. 

Phleum pratense. Scarce. Meadow and sunny orchard. 
Anthoxanthum odoratum. Common. Moist soil. Banks, orchard, ete. 
Digitaria Ischaemum. Common. Lawn, tennis court, ete. 

D. sanguinalis. Abundant. Lawn, garden, shrubberies, ete. 
Selaria glauca. Abundant. Meadow, garden weed. 

Carex spicata. Rather scarce. Orchard. 

Asparagus officinalis. Occasional. Dry upper orchard. 
Ornithogalum umbellatum. Frequent. Meadow, shrubberies, ete. 
Rumex Acetosa. Abundant. Rich soil in restricted area of orchard. 
R. Acetosella. Abundant. Sterile, acid soils throughout. 

R. crispus. Common. Meadow. 

R. obtusifolius. Scarce. Meadow. 

Polygonum aviculare. Abundant weed in thin lawn, sidewalks, ete. 
P. Convolvulus. carce and transient. Sunny bank of meadow. 
P. Persicaria. Common. Perennial beds, garden, meadow. 
Chenopodium lanceolatum. Frequent in orchard. 

Portulaca oleracea. Common garden and lawn weed. 

Stellaria media. Abundant weed in disturbed moist soils. 
Cerastium vulgatum. Abundant, chiefly as weed in lawn. 
Ranunculus repens. Common. Meadow, moist lawn, shrubberies. 
R. acris. Common. Drier portions of orchard. 

Berberis vulgaris. Frequent as seedlings under trees, shrubberies. 
Capsella Bursa-pastoris. Very scarce. Thin lawn. 

Barbarea vulgaris var. arcuata. Very scarce. 

Sedum purpureum. Abundant and aggressive. 

Trifolium agrarium. Very scarce. Dry sandy soil. Orchard. 

T. pratense. Scarce. Moist rich slope. Orchard. 

T. repens. Abundant, moist sunny orchard. Lawn. 

Vicia Cracca. Abundant in two areas of orchard. 

Rhamnus Frangula. Sporadic as seedling in strawberry bed. 
Daucus Carota. Abundant. Meadow, orchard, etc. 


66 Rhodora [ Vor. 60 


Lysimachia Nummularia. A single large patch near the White Pine. 

Prunella vulgaris. Abundant. Moist lawn, meadow, etc. 

Solanum Dulcamara. Common. Moist ground under shrubberies, trees, etc. 
Verbascum Thapsus. Frequent as seedlings in light soil. 

Veronica serpyllifolia. Abundant, moist lawn. 

Plantago lanceolata. Common. Lawns, meadow, ete. 

P. major. Abundant except in orchard. 

X Lonicera bella. Sporadic as seedlings under apple trees. 

Achillea Millefolium. Scarce. Dry, sunny orchard. 

Chrysanthemum Leucanthemum var. pinnatifidum. carce. Edge of meadow. 
Tanacetum vulgare. Sporadic. Nursery. 

Arctium minus. Sporadic as seedlings. 

Cirsium vulgare. Sporadic as seedlings. 

Cichorium Intybus. Scarce. Meadow, moist orchard. 

Leontodon autumnalis. Abundant. Meadow, orchard, lawn. 

Taraxacum erythrospermum. Formerly abundant in drier parts of lawn. 

T. officinale. Formerly abundant in meadow and richer portions of lawn. 
Hieracium aurantiacum. Scarce. First noticed in 1955. 

H. florentinum. Very scarce. Dryish ground in orchard. 


This list totals 60 species in 45 genera from 21 families. 


Those species of the foregoing genera which more or less resist 
an attempt to eradicate them appear to be assisted in their 
continued occurrence by the following agencies: Daucus, ger- 
mination of long dormant seeds(?); Solanum, birds; Arctium, 
animals; Cirsium, wind; Taraxacum, wind; Hieracium aurantiacum, 
wind. Because of their customary appearance as_ seedlings 
under trees and food-yielding shrubs the following doubtless are 
also spread by birds: Asparagus, Berberis, Lonicera. I am at a 
loss to account for the prevalence of Sedum purpureum and of its 
constant appearance as a weed in the infrequently cultivated por- 
tions of the garden, such as the strawberry bed. 


TABLE 2 
SPECIES INTRODUCED OR ADVENTIVE AND NATURALIZED FROM THE NEW WORLD 


Mollugo verticillata. Common weed. 

Geranium Robertianum. Abundant. Weedy in shrubberies and elsewhere. 

Cuscuta campestris. A single vigorous plant appeared in 1957, parasitic on 
garden carrot. 

Solanum rostratum. A single plant, 1956. Garden. 

Aster novae-angliae. Scarce. Meadow. First appeared about 1953, pos- 
sibly locally native. 

Galinsoga ciliata. Occasional. Garden and disturbed ground. 

Helianthus tuberosa. Meadow. Resists eradication. 

Rudbeckia serotina. Abundant. Orchard and dry banks. 


This list totals 8 species in 8 genera from 5 families. 


1958] Eaton,—Spontaneous Flora in Eastern Mass. 67 


TABLE 3 
NATIVE IN EASTERN MASSACHUSETTS 
BUT PRESENT POPULATIONS PROBABLY ORIGINATING FROM OUTSIDE THE LOT 


Pinus Strobus. Occasional as seedlings along walls. 

Festuca rubra. Common. Orchard. 

F. rubra var. commutata. Apparently scarce. Unmowed bank. 

Poa pratensis. Common. Meadow, orchard. 

Eragrostis spectabilis. Frequent. Orchard. 

Agropyron repens var. subulatum. Abundant. Meadow, orchard. 

Agrostis perennans. Frequent. Orchard. 

Muhlenbergia frondosa. Frequent. Shrubberies, disturbed ground. 

M. mexicana. Scarce, several clumps. Orchard. 

M. Schreberi. Frequent. Shrubberies, disturbed ground. 

Paspalum ciliatifolium var. Muhlenbergii. Rather common. Orchard. 

Panicum capillare var. occidentale. Not common. Orchard. 

P. lanuginosum var. fasciculatum. Frequent. Orchard. 

P. lanuginosum var. implicatum. Frequent. Orchard. 

P. spretum. Scarce. Orchard. 

Echinochloa pungens var. Wiegandii. Scarce. Orchard. 

Andropogon scoparius var. frequens. Scarce. Orchard. 

Cyperus strigosus. Frequent. Orchard, meadow, cultivated ground. 

Carex annectens. Not common. Orchard. 

Juncus tenuis. Abundant weed. Tennis court. 

Maianthemum canadense. Abundant. Edge of meadow under white pine. 

Sisyrinchium atlanticum. | Scarce. Orchard. 

S. montanum var. cerebrum. Scarce. Orchard. 

Habenaria lacera. Two plants. Orchard. First observed in 1957 as new 
arrival(?). 

Spiranthes cernua. Sporadic. Orchard. 

Carya glabra. Frequent as seedlings. Orchard. Several young fruiting 
trees. : 

C. tomentosa (?). A single sapling about 8 years old. Bank, edge of meadow. 

Corylus americana. Scarce. Thicket around large boulder. 

Betula populifolia. Strawberry bed, as occasional seedlings. 

Quercus alba. Frequent as seedlings especially under apple trees. Orchard. 

Q. rubra. Frequent as seedlings. Orchard. 

Ulmus americana. Property line, 2 tall trees. Occasional as seedlings under 
shrubs. 

Pilea pumila. Abundant weed in one location under shrubs. 

Polygonum H ydropipiper. Weed in vineyard, perennial bed. 

P. pensylvanicum var. laevigatum. Weed in vineyard, perennial bed. 

Phytolacca americana. Common weed in shrubberies, ete. 

Sagina procumbens. Common weed along sandy flagstone path, shrubbery. 

Spiraea latifolia. Very scarce but surviving repeated burns. Orchard. 

Fragaria virginiana. Two large clones. Orchard. 

Potentilla canadensis. Common. Orchard, thin lawn. 

Geum laciniatum var. trichocarpum. Scarce. Plantation, shrubberies. 

Rubus Enslenii. Very scarce. Orchard. 

R. occidentalis. Frequent as seedlings under old trees, along walls, ete. 

R. pensylvanicus. Encroaching from neighboring field. 

Prunus serotina. One well grown tree. Frequent as young sprouts in 
orchard. 

P. virginiana. Several mature specimens. Thicket. 


68 Rhodora [Vor. 60 


Ozalis europaea forma villicaulis. Abundant. Orchard, lawn, shrubberies. 

Geranium maculatum. Well established near plantation. 

Acalypha rhomboidea. Common. Orchard, moist shrubberies, garden. 

Euphorbia supina. Abundant weed. Driveway, thin lawn. 

Celastrus scandens. A single old vine. Numerous root shoots in vicinity. 

Impatiens capensis. Abundant. Edge of meadow. 

Parthenocissus quinquefolia. Orchard, along southwall. Also as frequent 
seedlings. 

Vitis aestivalis. A single specimen, recently established. Shrubbery. 

Hypericum mutilum var. parviflorum. Frequent. Moist open shrubbery. 

H. perforatum. Scarce. Orchard. 

Viola cucullata forma albiflora. Scarce. Meadow, orchard. 

Epilobium coloratum. Frequent. Moist open shrubbery. 

Circaea quadrisulcata var. canadensis. Frequent. Moist open shrubbery. 

Cornus alternifolia. Scarce. Orchard thicket. 

Vaccinium angustifolium var. laevifolium. Searce. Orchard, slope. 

Lysimachia quadrifolia. Large clone, under white pine. 

Fraxinus americana. Frequent. Shrubberies, etc., as seedlings. 

Asclepias syriaca. Nearly eradicated. Formerly abundant. 

Convolvulus sepium. Persistent in three locations. 

Verbena urticifolia. Scarce. Plantation. Sporadic elsewhere. 

Solanum americanum. Sporadie weed. Garden. 

S. carolinense. Large clone. Orchard. Spreads rapidly unless checked. 

Physalis heterophylla. Upper orchard. Thin sandy soil where under observa- 
tion since 1940. Two or three stems first appeared in 1956 under old 
apple tree north side of house on disturbed sandy bank. 

Linaria canadensis. Scarce. Shrubbery. 

Orobanche uniflora. Very scarce and sporadic. Edge of meadow (1946). 
Orchard near south wall (1952). 

Plantago Rugelii. Ubiquitous weed. Meadow, lawn, shrubberies, etc. 

Solidago caesia. Persistent in a single location behind tennis court. 

. canadensis. Frequent, formerly abundant. Meadow, lower orchard. 

S. juncea. Frequent. Orchard. 

. nemoralis. Frequent. Orchard. 

S. rugosa. Common. Orchard, meadow. 

Aster ericoides. Scarce. Orchard. 

A. lateriflorus. Common. Orchard. 

A. lateriflorus var. pendulus. carce. Orchard. 

A. linariifolius. Scarce. Orchard. 

A. undulatus. Abundant. | Orchard. 

Erigeron annuus. Abundant. Meadow, orchard, lawn. 

E. canadensis. Common. Orchard. 

E. pulchellus. Several large clones. Orchard. 

Antennaria neglecta. Common. Orchard. 

A. neodioica. Common. Orchard, lawn. 

A. plantaginifolia. Scarce. Orchard. 

Gnaphalium obtusifolium. Common. Orchard. 

Ambrosia artemisiifolia var. elatior. Abundant. Orchard. 

Bidens frondosa. Sporadic. Shrubbery. 

Lactuca canadensis var. latifolia. Meadow where weedy. Orchard where fre- 
quent. 

L. canadensis var. longifolia. Common. Orchard. 


Nn 


Nn 


This list totals 88 species in 65 genera from 37 families. 


1958] Eaton,—Spontaneous Flora in Eastern Mass. 69 


Assignments to the category of Table 3 are the result chiefly 
of subjective judgment. One basie assumption which has influ- 
enced my opinion is that the lot had been mowed annually for 
many years right up to the property lines, except for a small thicket 
near a big boulder in the northeast corner. The reasons for this 
assumption, not stated here, seem to me to be convincing. There- 
fore, all the spontaneous trees and most of the shrubs are listed 
here. Those native herbaceous plants collected in the unplowed 
orchard, which appear to require the specialized habitat of dry, 
sunny sterile soils, presumably have invaded the lot after it was 
originally cleared. Furthermore, it is a fact that the plantation 
and most of the shrubberies were established by me after 1940. 
There seems to be little doubt about the recent invasion of those 
species found only in the artificial habitats of plantation, shrub- 
beries, tennis court, lawns, driveways and other disturbed soils. 
It is conceivable, however, that Corylus, Prunus virginiana (both 
immediately adjacent to the boulder in the thicket), Geranium ma- 
culatum, Cornus, Asclepias, and Solanum carolinense have been 
present in the lot since pre-colonial times. 

There are several species listed in Table 3 which appear to occur 
only in disturbed or artificial soils, in the leaf mold of shrubberies, 
or in mulched areas where there is little competiton from aggres- 
sive herbaceous vegetation. These, referred to by generic name 
only in cases where the identities are in no doubt, are as follows: 
Muhlenbergia (both species), Juncus, Betula, Ulmus (the seedlings), 
Pilea, Polygonum (both species), Phytolacca, Sagina, Geum, 
Geranium, Euphorbia, Impatiens, Vitis, Hypericum, Epilobium, 
Circaea, Fraxinus, Verbena, Solanum americanum, Linaria, Bidens. 


TABLE 4 


NATIVE IN EASTERN MASSACHUSETTS 
AND VERY POSSIBLY PERSISTENT 77 situ, FROM PRE-COLONIAL TIMES 


Equisetum arvense. Abundant. Meadow, borders, garden weed. 

Osmunda cinnamomea. carce. Orchard, chiefly along south wall. 

O. Claytoniana. Common. Orchard, chiefly along south wall. 

O. regalis var. spectabilis. Very scarce. Orchard, in moist sod land. 

Onoclea sensibilis. Abundant. Meadow, orchard. 

Dryopteris Thelypteris var. pubescens. Abundant. Orchard. 

Athyrium Filix-femina var. Michauzii. Frequent. Orchard, along south wall. 

A. Filir-femina var. Michaurii f. elatius. Frequent. Orchard, along south 
wall. 

A. Filiz-femina var. Michauxii f. laurentianum. Very scarce. Orchard. 

A. thelypterioides f. acrostichoides. Very scarce. Orchard. 


70 Rhodora | VoL. 60 


Carex cephalophora. Frequent. Orchard. 

C. normalis. Rather scarce. Orchard. 

C. pallescens var. neogaea. Rather scarce. Orchard. 

C. pensylvanica. Frequent. Orchard. 

C. scoparia. Not common. Orchard. 

7, Swanti. Very scarce. Orchard. 

C. tenera. Common. Orchard. 

Luzula multiflora. Rather common. Orchard. 

Apios americana. Two colonies. Along north and south walls. 

Rhus radicans. Abundant and ubiquitous. Orchard. Seedlings repeatedly 
appearing in shrubberies, under trees, nursery, etc. 

Viola papilionacea. Abundant. Orchard. Also as weed in shrubberies. 

Apocynum androsaemifolium. A single large clone. Orchard near thicket. 
Also one seedling found in raspberry bed in 1956. 

Aster cordifolius. Common. Orchard. Along walls. Also abundant weed 
in shrubberies and disturbed ground. 

A. novi-belgii. Scarce, nearly extirpated. Meadow. 

A. puniceus var. compactus. Scarce, nearly extirpated. Meadow. 

A. vimineus. Common. Orchard. 


This list totals 24 species in 12 genera from 10 families. 

Many of the species which I have assigned to Table 3 possibly 
were sparse or rare constituents of the original flora of the lot. 
However, I have excluded from Table 4 most of the forage plants 
(those grasses, sedges and other herbaceous species frequently dis- 
seminated directly or indirectly by grazing animals), and also 
aggressive species shedding wind-borne seeds or attractive to 
seed- and fruit-eating birds. The mere fact that with due diligence 
we can find most of them in our fourth-growth woodlands is no 
proof that they occurred normally in a virgin deciduous forest 
except as transients in naturally disturbed soils or habitats, such 
as blow-downs and “burns” started by Indians or rarely by light- 
ning. These considerations lead to the inevitable reflections con- 
cerning the profound change in the composition and relative 
abundance of our local flora, which has occurred during the past 
three hundred years. The conversion of the wilderness to farm 
land with its multiplicity of wood lots, pastures, mowings, fence 
rows, stone walls, ditches, roadsides and thickets offers numerous 
suitable habitats for hundreds of species where few or none existed 
under primitive conditions. Hence, a large number of our native 
species, now common to abundant throughout the countryside, 
originally must have been uncommon, or local, or even rare, for 
the simple reason that favorable habitats were scarce. 

In attempting to sort out the species which may have survived 
in situ as a relic of the primeval flora of the lot it is first necessary 
to reconstruct the probable nature of the forest growth before the 


1958] Eaton,—Spontaneous Flora in Eastern Mass. 71 


land was cleared and to visualize the various ecological niches pro- 
vided by it, with due reference to physical characteristics of the 
site as described earlier in this paper. It is then possible to con- 
sider whether the ecological requirement of each native species now 
growing on the lot is consistent with the ecological niches reason- 
ably assumed to have been provided by its original forest cover. 
As a clue to this problem, there is a patch of rocky woods on the 
steeper north slope of our hill at a distance of about three hundred 
meters but at a somewhat lower elevation. Although badly mauled 
by the series of hurricanes during the past twenty years, it has 
every appearance of being a relatively undisturbed remnant of the 
original forest. It consists mainly of a mixed deciduous hardwood 
forest. A brief winter inspection reveals the following species 
(doubtless among others): Pinus rigida, Juglans cinerea (a single 
aged and dying specimen), Carya sp., Betula lenta (now the most 
abundant species and rapidly invading the openings made by 
windfalls), Quercus alba, Q. rubra, Ulmus americana and Fraxinus 
americana. There are also a few large stumps of Castanea dentata. 
At the lower edge of this wooded patch is a small water course 
which, in fact, is a continuation of the covered drainage ditch in 
my meadow. Its herbaceous vegetation is what one might expect 
in an area such as this: ferns, grasses, sedges, and the other ele- 
ments of a flora to be found in dry to moist woods, glades, and 
along partially shaded runnels. Prior to the hurricanes there were 
several very large old white oaks on the southern flank of our hill, 
and, according to an old neighbor, a small grove of ancient chestnut 
trees in the vicinity, most of which were removed between 1875 
and 1900. 

It is probable, therefore, that mature deciduous trees grew orig- 
inally on my portion of the hill, effectively suppressing sapling 
growth under them and offering a suitable habitat for a varied 
herbaceous flora in dry, moist and swampy open woods. I con- 
clude that the foregoing list represents a remnant of the original 
flora—a remnant capable of persisting under the harsh ecological 
conditions imposed by mowing, burning and competition with in- 
troduced species. 

It is of some interest, perhaps, to note that the following plants 
listed in Tables 3 and 4, although indigenous to the lot or in the 
neighboring region, behave in a weedy manner in one or more of the 
various "artificial" habitats on my place, viz., meadow, garden, 


72 Rhodora [ Vor. 60 


shrubberies, plantation, lawn, tennis court, but occur sparingly if 
at all on undisturbed soils: Equisetum, Onoclea, Agropyron, Muh- 
lenbergia sp., Cyperus, Juncus, Pilea, Polygonum sp., Phytolacca, 
Sagina, Geum, Potentilla, Geranium, Oxalis, Acalypha, Hypericum 
mutilum, Epilobium, Circaea, Plantago ssp., Aster cordifolius, 
Erigeron annuus, Helianthus, Bidens, Lactuca. 

In certain instances there is indirect evidence of the means of 
dissemination, aside from those plants whose seeds are obviously 
windblown or have been introduced in manure and lawn grass 
seed. Seedlings of the following plants occur almost exclusively 
under apple trees used by nesting robins or under juicy fruited 
shrubs attractive to catbirds, starlings, and robins, and other fruit- 
eating birds: Asparagus, Phytolacca, Berberis, Rhus, Partheno- 
cissus, Rhamnus (strawberry bed), Physalis, Lonicera. I have 
alluded to the surprising appearance of Vitis aestivalis. Mention- 
ing the occurrence to an ornithological friend, he asked if I had 
ever seen Ruffed Grouse on my place. When I assured him that 
I had recorded Grouse on at least three widely separated occasions 
he said that Grouse were fond of ripe grapes and apparently ate 
them whole. I have never permitted Arctium or Bidens to reach 
maturity on my place, but having observed that my dog frequently 
comes home with an assortment of burs in his coat, it is reasonable 
to assume that long coated animals (dogs, cats, skunks or foxes) 
have transported the seeds in these two cases. The frequent oc- 
currence of hickory and oak seedlings in the orchard is probably 
due to Gray or Red Squirrels, although Jays and Crows cannot be 
excluded as possibilities. 

The Flora of the Boston District, published serially in RHODORA, 


TABLE 5 


AGGREGATE COUNT OF FAMILIES, GENERA AND SPECIES COMPARED WITH 
THE FLORAS OF THE BOSTON DISTRICT AND OF GRAY'S MANUAL RANGE 


"EN 
Species 
Families Genera — —— —— 
Introd. |Native| Total | % Native 
7C 


Lincoln House Lot 53 95 66! 


110? 176 62.5 


Boston District 133 697 7890! | 1312? | 2101 62.5 
Gray's Manual Range 168 | 1133 1098 | 4425 | 5523 80.1 


! Introduced and naturalized or adventive from outside eastern Massachusetts. 
? Native in eastern Massachusetts. 


1958] Jennings,—Thelepogon elegans Roth 73 


Vols. 9-26 incl., comprises a preliminary list of all Pteridophytes 
and Spermatophytes, including varieties and named forms, re- 
liably reported to a committee of the New England Botanical 
Club as growing without cultivation in a precisely defined area 
which is here roughly described as within thirty miles of Boston. 
Lincoln is fifteen miles due west of Boston. 

The Boston District figures are derived from an unverified count 
of the taxa involved. Accuracy, as of the present day, is impossible 
because the list has never been brought up to date either in respect 
to nomenclature or additions. However, it is unlikely (by the law 
of chances) that the ratio of native species to total species would be 
significantly different were an up-to-date list available. Inci- 
dentally, it is, of course, a sheer coincidence that this ratio of 
62.5%, as stated above, is exactly the same as that for the Lincoln 
house lot! The significant point is that they are of the same order 
of magnitude and are in marked contrast to the ratio of 80.1% for 
Gray's Manual range. Here, it seems, is a bit of concrete evidence 
of the truth of the commonly accepted assumption that the exotic 
elements of the floras of long and densely settled areas of this 
country (e.g., along the eastern seaboard) are much larger than of 
the relatively youthful regions (e.g., Mississippi basin and prairie 
states). It takes time for introductions to spread, even when 
suitable habitats are available. Perhaps in this age of universal 
motor travel the tempo of the spread is being greatly accelerated. 

The figures in Table 5 well illustrate the diversity of our local 
flora. My house lot of less than three acres contains nearly 8.5% 
of the species reported from the Boston District (approximately 
1900 square miles of which a substantial fraction is salt marsh and 
sand dune). The Boston District, less than one-quarter of the area 
of Massachusetts, contains 38% of the species recorded from the 
entire Gray's Manual range. This diversity is even more striking 
in respect to genera and families.—LINCOLN, MASSACHUSETTS. 


THELEPOGON ELEGANS ROTH IN THE DOMINICAN REPUBLIC.— 
Specimens of this tropical Old World grass recently came to hand 
for determination, with the information that it is becoming a pest 
in the rice fields of the Dominican Republic. In its immature 
stages it resembles rice plants rather closely, thus making more 
difficult the application Fof selective control measures.—O. E. 
JENNINGS, CARNEGIE MUSEUM. 


74 Rhodora [Vor. 60 


THE FRUTICOSE AND FOLIOSE LICHENS OF 
WORCESTER COUNTY, MASSACHUSETTS. 


VERNON AHMADJIAN 


The study of lichens has undoubtedly been discouraging to many 
who have become interested in these forms, due to the lack of sim- 
ple and readily available keys. It is hoped that this work will be 
useful in the identification of lichens and help to contribute to the 
fulfillment of this major need in the field of lichenology. 

Although the present work is regional in its scope, the lichens 
described are by no means localized and can be readily found 
throughout the northeastern part of the United States. The keys, 
however, have been designed specifically from material collected 
Within Worcester County. Anyone attempting to determine 
lichens from outside of this region should bear this in mind. 

Only the fruticose and foliose lichens have been dealt with be- 
'ause the erustose forms are too difficult to be given a simple and 
yet adequate treatment suitable for beginners. 

Worcester County, making up the central part of Massachusetts, 
is the largest county in the state, and consists of an area of approxi- 
mately 1,522 square miles or 974,000 acres. It extends 35 miles 
from east to west and 50 miles, or the entire width of the state, 
from north to south. The county is characterized by ranges in 
elevation of from 800-1,200 feet, excluding the extremely low and 
high points, and shows a remarkable uniformity of the horizon line. 
The most notable of the remnants of elevations above the plateau 
surface still existing are Mt. Wachusett in Princeton, elevation 
2018 ft., Little Wachusett, Princeton, elevation 1,559 ft. and 
Asnebumskit Hill, Paxton, elevation 1,395 ft. Throughout the 
county many rocks have been scattered as a result of the glacia- 
tion; contacts of schist and granite are most prevalent. 

The climate of Worcester County is as variable as that in any 
temperate region. In general, the winter temperature averages 
about 26 degrees with a minimum recorded of — 24 degrees and the 
average summer temperature is about 69 degrees, with a maximum 
recorded of 103 degrees. The average annual precipitation is 42.3 
inches. 


!'This contribution is part of a thesis submitted to Clark University, Worcester, Massa- 
chusetts, for the degree of Master of Arts, 1954-1956. 


1958] Ahmadjian,—Frutieose and Foliose Lichens 15 


ACKNOWLEDGEMENTS 


I wish to express my gratitude to Dr. David Potter, Chairman of the De- 
partment of Biology, Clark University, and Dr. I. Mackenzie Lamb, Director 
of the Farlow Herbarium, Harvard University. 

I wish also to express my thanks to the following people who gave willingly 
of their help: Dr. Grace E. Howard, Dr. Alexander W. Evans, Dr. Mason E. 
Hale, Dr. John W. Thomson, Dr. Henry A. Imshaug, Dr. Burton N. Gates, 
Dr. Stuart K. Harris, and Miss June M. Allstrom. 


LITERATURE USED IN THE IDENTIFICATION OF LICHENS 


For an introduction to lichenology, the Lichen Book by G. G. 
Nearing has proved to be most satisfactory. It describes approxi- 
mately 500 lichens, most of which are commonly found in the north- 
eastern part of the United States, and its illustrations and descrip- 
tions are good. Unfortunately, the approach is rather unscientific 
and the keys are not too usable. 

Lichens by Annie Lorrain Smith is an excellent book embracing 
virtually all aspects of lichenology, from the history of lichens to 
their physiology, ecology, morphology, ete. It also has an exten- 
sive bibliography and is an invaluable book for those who are 
interested in studying this group of plants. 

The Lichen Flora of the United States by Bruce Fink gives a 
comprehensive survey of the lichens of the United States and serves 
as a good reference book. Its keys, however, are difficult and 
poorly constructed and on the whole the beginner would find the 
book rather cumbersome. 

For those wishing to go on further in the study of lichens, Raben- 
horst Kryptogamen Flora provides an excellent treatment, with 
keys and descriptions of the various lichen groups. Each group 
treatment is the work of an expert in that particular area. "The 
section on lichens in Engler and Prantl, Die Natürlichen Pflanzen- 
familien, is also worthy of notice here, containing good keys and 
descriptions. 

The Catalogus Lichenum Universalis by A. Zahlbruckner is 
another indispensable piece of lichenological literature containing 
all known lichen taxa up to 1940, and giving nomenclature and 
references to literature pertaining to each lichen that is listed. 
Consisting of ten volumes, it contains over 17,000 lichen names, 
many of which, however, have now been outdated. Dr. I. Mac- 
kenzie Lamb is at present working on an extension of this treatment 
by Zahlbruckner which will bring this list of known lichens up to 
date. 


76 Rhodora [ Vor. 60 


The Lichen Handbook? by Mason E. Hale is an excellent up- 
to-date booklet on lichenology and includes sections on the chem- 
istry, reproduction, phytogeography, and classification of lichens. 
Prepared from a series of lecture notes on the subject, it reviews all 
aspects of the field and would be of much value to beginners. 

For recent lichenological works or studies, The Bryologist, : 
quarterly journal of the American Bryological Society, would be a 
likely source containing a section listing recent literature on lichens 
by William L. Culberson. Rhodora, a monthly journal of the 
New England Botanical Club, is another possible reference source. 


= 


KEY TO THE Matin Groups or LICHENS 


Thallus elongated, shrubby to band-like or thread-like, erect or pendulous, 
usually branched; arising either directly from the substratum by a basal 
holdfast, or from a primary thallus, in some instances dying off at the 
base... lesse la Fruticose Lichens. 

Thallus flattened, leaf-like, dorsiventral, with a visible difference between 
upper and under surfaces; partly free from the substratum, to which it is 
relatively loosely attached, usually by rhizoids......... Foliose Lichens. 

Thallus crust-like, closely applied and wholly fused to the substratum by the 
underside; not detachable except by scraping off in small fragments; 
commonly areolate, divided by cracks into small island-like portions 

Crustose Lichens. 


KEY TO THE GENERA OF FRUTICOSE LICHENS 


1. Thallus flattened; primary thallus absent... 2. ....00000.00 00000000005. 2 

1. Thallus not flattened, cylindrical; primary thallus present or absent. .... 4 

2. Thallus soft and flaccid when dry, usually pendulous. . . £vernia mesomorpha 
Nyl. On bark. "Thallus greenish or yellowish-green, sorediose. Spores 
simple, one-celled. 

2. Thallus rather stiff and rigid, erect... .....00000.00000 00 cece eee eee 3 

3. Thallus yellowish, almost always without marginal spinules...... Ramalina 
pollinaria (Liljebl.) Ach. Branches short, tips bursting open, densely 
farinose sorediate. Spores two-celled. 


3. Thallus brownish, with marginal spinules............... CETRARIA, p. 77 
4. Stalks solid... iilii iiie eee eee 5 
4. Stalks hollow or loosely filled........ liliis 7 
5. Stalks with tough tensile cartilaginous central core... .........0.0..00% 6 
5. Stalks without tough tensile cartilaginous central core... .Baeomyces roseus 


Pers. On sandy banks along roadside. Very common. Primary 
thallus erustose, gray. Podetia simple, white or pinkish, bearing single 
mushroom-like pink apothecia. Thallus bearing scattered white or 
greenish globules. 
6. Thallus greenish to silver-gray; assimilative branches squamulose, coralloid, 
or granulose, n... naunan 000 eee STEREOCAULON, p. 81 
6. Thallus greenish-gray; assimilative branches filamentous. ... . USNEA, p. 81 


? Available from the author: Mason E. Hale, Division of Cryptogams, Smithsonian Institu- 
tion, United States National Museum, Washington 25, D. C. 


1958] Ahmadjian,—Frutieose and Foliose Lichens 77 


7. Thallus hair-like...... Alectoria nidulifera Norrl. On rocks and rotting 
wood. "Thallus sorediate. Apothecia rare. 
7. Thallus cylindrical, shrubby or dendroid................ CLADONIA, p. 77 


IKEY TO THE SPEcrES or FnvTICOsE LICHENS 
CETRARIA 


1. Thallus having scattered brown or black tubercles. Apothecia originating 
from under surface................... Cetraria ciliaris Ach. On bark. 
1. Thallus having no tubercles. Apothecia, if present, originating from upper 
surface...... Cetraria islandica (L.) Ach. On ground in pine woods. 
Stalks strap-shaped, forking, thin, curling when dry to form a trough-like 
structure. 
CLADONIA 


1. Primary thallus crustose, gray. Podetia simple or branched, brown 
tipped. Apothecia brown and clustered. . . . Cladonia papillaria (Ehrh.) 
Hoffm. On ground. 


1. Primary thallus squamulose or disappearing.................0. 00.000 2 
2. Primary thallus disappearing, rarely present. Podetia branched, usually 

elongated and entangled, dying off at the base..................... 3 
2. Primary thallus squamulose, persistent. ............ sse 12 


3. Podetia squamulose, usually erect and sterile, showing openings in branch 
axils, grayish-green, rarely brownish. Apothecia, if present, brown. . .4 

J. Podetoa Te NINNUDIDIOSB es vs sins a a.) VINEIS. 5 

4. Podetia slender, entangled, warty in appearance. KOH —, Pd+ red 
Cladonia furcata (Huds.) Schrad. On ground. Var. palamaea (Ach.) 
Vainio. Podetia brownish. Var. pinnata (Flk.) Vainio. Podetia 
densely squamulose. 

4. Podetia somewhat swollen, little entangled, covered sparingly with 
granular soredia. KOH —, Pd+ red....Cladonia scabriuscula (Del.) 
Vainio. On ground. 

5. Podetia entangled, short, spinose at tips, usually forming sponge-shaped 


tws OH ie a ee ee ee 6 
5. Podetia entangled, or little entangled. Extreme tips blunt to hair-like, 
BEND OL erecta. man ihe ek, fot wes ete es 7 


6. Extreme tips spinose, branch axils usually perforated. . . Cladonia uncialis 
(L.) Web. On ground in rocky hillside pastures. Very common. 

6. Podetia much enlarged, contorted, flabby, showing lattice-like openings 

Cladonia Boryi Tuck. On ground. 

7. Podetia abundantly farinose-sorediate, little entangled. KOH—, Pd+ 

c ra Cladonia farinaceae (Vainio) Evans. On ground. 


~ 


8. Podetia silver-gray and smooth or warty to granular in older specimens. 
Tips brownish, usually divided into whorl-like arrangements and then 
each tip subdivided into many drooping finger-like projections. KOH 
+ yellow, Pd + red...... Cladonia rangiferina (L.) Web. On ground in 
rocky hillside pastures. Fairly common. f. crispata Coem. Podetia 
more densely branched and colonies more compact. 

8. Podetia greenish or yellowish-gray................ llle cee eee 9 

9. Podetia slender, tips forked, hair-like; branch axils usually closed. KOH 
+ yellow, Pd + red..... Cladonia tenuis (Flk.) Harm. On ground in 
open fields and hilly pastures. Very common. 


Rhodora [ Vor. 60 


Podetia polytomously branched; branch axils usually perforated... ... 10 

Tips not usually darkened, tending to curve in one direction. KOH —, 
Pd + red...... Cladonia sylvatica (L.) Hoffm. On ground in rocky 
hillside pastures. 


. KOH —, Pd —.............susssseseseee ha 11 
. Podetia forming rather loose branch-systems; extreme tips rather long, 


diverging in various directions. ...Cladonia mitis Sandst. On ground. 


. Podetia thickened, tips short and numerous. . . . Cladonia submitis Evans. 


On ground. 


. Podetia usually lacking or very short. Apothecia, if present, brown, 


resting on or appearing immersed in the primary squamules........ 13 


. Podetia present, well-developed, simple or branched, cylindrical or cup- 


forming... oona auauua eras 14 


3. Primary squamules small, margins serrated into fine tooth-like lobes. 


Usually sterile. Apothecia, if present, large, lumpy. KOH —, Pd + 
red...... Cladonia caespiticia (Pers.) Flk. On ground, rotting wood, 
bark. Very common. 


. Primary squamules long, strap-shaped, notched at tips, blue-gray, curling 


upward when dry showing white under surface. KOH + yellow, Pd + 
red... 2.2.0... Cladonia apodocarpa Robbins. On ground. Common. 


. Podetia clustered to form more or less matted colonies; apothecia brown. 15 
. Podetia separate, not forming mat-like colonies..................... 18 
. Podetia eupless, having openings in branch axils; often squamulose. KOH 


—, Pd + red... onana aaa aaa aaa Cladonia furcata (Huds.) Schrad. 


. Podetia usually covered in various degrees with large or minute squamules; 


usually forming open cups which broaden out gradually or abruptly: 


irregular dichotomous branching... .......0.0.0.00.00000 000 cece eee 16 
. Podetia forming cups with sieve-like openings. . . . Cladonia turgida (Ehrh.) 
Hoffm., f. squamulosa (Rabenh.) Theobald. On ground. Podetia 
squamulose. 
. Podetia not forming cups with sieve-like openings. ................... 17 
. Podetia densely covered with minute or large squamules. KOH —, 


Pd —....Cladonia squamosa (Scop.) Hoffm. f. squamosissima Flk. 
Podetia densely squamulose. 


7. Podetia not densely covered with squamules; forming well developed cups 


2-4 mm. in diameter. KOH +, Pd +....Cladonia carassensis Vainio, 
f. regularis Vainio. On ground. Podetia repeatedly proliferating with 
short stories (2-13 mm.) Cups small, regular, usually abruptly dilated. 
No squamules. 


. Podetia seyphous, forming cups... isses 19 
. Podetia ascyphous, not cup-forming.........i isse eee 34 
. Podetia cylindrical, except at the extreme tips where they expand abruptly 

to form small cups. Apothecia brown.........sse esee eene 20 
. Podetia expanding gradually to form rather large cups............... 25 
. Cups funnel-shaped, usually opening into podetial cavity. Podetia never 

sorediose..... clesie eme 21 
. Cups closed, shallow. Podetia sorediose......... llle. 22 
. KOH —, Pd —...........0..0...... Cladonia squamosa (Scop.) Hoffm. 
. KOH +, Pd +... aaa Cladonia carassensis Vainio. 


. Podetia farinose-sorediate throughout except for a small portion at base. 


Cups merely small depressions in slightly expanded tips, often lacking 
and the podetia pointed and horn-like at the tips. KOH —, Pd + 


1958] Ahmadjian,—Fruticose and Foliose Lichens 79 


32. 


33. 


red. ...Cladonia coniocraea (FIk.) Spreng. On ground, rotting wood, 
bark. Very common. f. phyllostrota (Flk.) Vainio. Podetia squamu- 
lose. f. stenoscypha (Stuckenberg) Sandst. Podetia forming cups, 
narrow at the mouth, often abortive. 


. Podetia granular-sorediate, or if farinose, mixed with granules. .......23 
. Podetia usually over 15 mm. long, covered with finely granular soredia 


only on upper part; expanded at tips to form shallow star- or wand- 
shaped cups; or cupless and simple. KOH —, Pd + red..... Cladonia 
nemoxyna (Ach.) Nyl. On ground and rotting wood. Very common. 


. Podetia short, less than 15 mm. long. KOH —, Pd + red........... 24 
. Podetia covered with farinose soredia, mixed with coarse granules. Cladonia 


cylindrica Evans. On ground. f. scyphifera Evans. Podetia stout, 
forming distinct cups at the tips. 


. Podetia covered with coarse granular soredia....Cladonia pityrea (L.) 
Hoffm. On rotting wood. Common. 
. Cups flat, shallow, saucer shaped, usually proliferating once or several 


ames. Apothecia brown... ae sd luii enr roro E: 26 
. Cups deep, goblet shaped, rarely proliferating, and then usually only 
once. Apothevia brown orred, ......,.. 55S esL. 27 
. Proliferations few, from margins of cups... .Cladonia gracilis (L.) Willd. 
On ground. 


. Repeated proliferations from center and margins of cups, or from sides of 


podetia. KOH —, Pd -F....Cladonia verticillata Hoffm. On rotting 
wood and ground. Very common. f. aggregata (Del.) Oliv. Pro- 
liferations central and numerous. f. apoticta (Ach.) Vainio. Podetia 
with proliferations along the sides as well as from the centers of the cups. 
f. phyllocephala (Flat.) Oliv. Podetia squamulose. 


. Apothecia, if present, red. Podetia pale bluish to yellowish-green....28 
. Apothecia, if present, brown. Podetia grayish-green to brownish..... 29 
. Podetia usually covered with coarse granular soredia, rarely farinose. 


KOH -, Pd —....Cladonia pleurota (L.) Willd. On ground. Com- 
mon. Var. frondescens (Nyl.) Oliv. Podetia squamulose. 


. Podetia dusted with fine soredia, farinose; long, usually contorted and 


stout. Proliferations, if present, from margin of cup. KOH —, 
Pug wes d WC Cladonia deformis (L.) Hoffm. On ground. 


9. Podetia esorediose. KOH —, Pd + red... .Cladonia pyzidata (L.) Hoffm. 


On ground and rotting wood. Very common. 


e sse eni cut es a Lane kee nea w'e esd Dae Cok DLN bebe as 30 
HIMOPPHIRTAEIIOSB elsi ive ce tnn hm yRRH iIIa ER» eles mote OE 
SISOIediaAJgrAnulBr. 6o REVUES VV wer eU eri ace ee eee 
. Cups narrow, slender, sometimes lacking, KOH —, Pd + red... . Cladonia 

fimbriata (L.) Fr. Soredia throughout length of stalks. On ground. 
. Cups rather well formed, rarely lacking............................82 
. Podetia forming distinct cups with marginal proliferations. KOH —, 


Pd + red... .Cladonia cornutoradiata Sandst. On ground. f. radiata 
(Schreb.) Sandst. Podetia forming distinct cups with marginal pro- 
liferations. 

Podetia forming broad, funnel-shaped cups. KOH + brown, Pd + 
pus Cladonia conista (Ach.) Robbins. Soredia found only on cups. 
On ground. 

KOH —, Pd + red....Cladonia chlorophaea (Flk.) Spreng. On ground 
and rotting wood. Very common. f. carpophora (Flk.) Anders. 


80 


33. 


34. 
34. 
35. 


35. 
36. 


36. 
37. 


37 
38. 


38. 
39. 


40. 


42. 


Rhodors [ Vor. 60 


Podetia fertile. Apothecia usually borne on proliferations from margins 
of cups. 

KOH —, Pd —....Cladonia Grayi Merrill. On ground and rotting wood. 
Very common. f. carpophora Evans. Podetia fertile. Apothecia 
usually borne on proliferations from margins of cups. f. prolifera 
Sandst. Cup-forming proliferations from margins of primary cups. 
f. squamulosa Sandst. Podetia squamulose. 


Apothecia red... e 35 
Apothecia brown... 20... .00.00 000000 cc es 40) 
Podetia esorediose, simple or branched, with or without squamules. 

Apothecia almost always present....Cladonia cristatella Tuck. On 


ground, rotting wood and bark. Very common. f. ochrocarpia Tuck. 
Apothecia yellow, to flesh colored. f. simulata Robbins. Bearing 
minute, clustered, pale apothecia on short lateral outgrowths of main 
podetial axes. f. squamosissima Robbins. Apothecia red.  Podetia 
densely squamulose. f. vestita Tuck. Apothecia red. Podetia squa- 
mulose. 


Podetia sorediose. 2... rre 36 

Thallus yellowish-green. Primary squamules densely farinose-sorediose, 
often crust-like. . . . Cladonia incrassata Flk. On rotting wood. Fairly 
common. 

Thallus greenish gray or whitish. ....... uao aaa aaa e eese 37 


Podetia densely covered with coarse granules and squamules, usually 
falling away at tips to expose whitish medulla. KOH —, Pd —, or 
Pd + pale yellow... . Cladonia didyma (Fée) Vainio. On rotting wood. 

Podetia covered with farinose, rarely granular soredia............... 38 

KOH + yellow, Pd + red....Cladonia macilenta Hoffm. On rotting 
wood and ground. Common. 


KOH —, Pd =... 39 
Podetia usually decorticate and farinose sorediate throughout length 
except small basal portion, slender... .Cladonia bacillaris (Ach.) Nyl. 


On rotting wood and ground. Very common. f. clavata (Ach.) Vainio. 
Forms with simple podetia which are blunt at tips; usually sterile. 
Apothecia, if present, always terminal. f. reagens Evans.  Podetia 
with yellowish-brown spots. KOH +, purple red. 

Podetia not decorticate throughout; farinose sorediate in upper 
parts....Cladonia Floerkeana (Fr.) Flik. On rotting wood and ground. 
J. trachypoda (Nyl.) Vainio. Podetia large, well developed, bearing 
cups; partly decorticate. 

Primary squamules containing large granules along the margins, forming a 
dense, granular crust. Podetia granulose sorediate. Thallus small, 


delieate....Cladonia. delicata. (Ehrh.) Fik. On rotting wood. Fairly 
common. 
Primary squamules not forming a granular crust). .0....0000.00..... 41 
Primary squamules small, usually 1 mm. long or less.......000000... 42 
Primary squamules large... oleo eese 43 


Podetia small, simple or branched sparingly at tips, usually contorted. 
Apothecia large, turban-like, dark brown. KOH + yellow, Pd + 
red. ...Cladonia capitata (Michx.) Spreng. (Syn. C. mitrula Tuck.) 
On ground. Fairly common. 

Podetia larger, often squamulose, producing short branches at tip. Apo- 
thecia large, buff or lavender....Cladonia piedmontensis Merrill, f. 
squamulosa Robbins. On ground. Podetia squamulosc. 


1958] Ahmadjian,—Fruticose and Foliose Lichens 81 


43. KOH + red, Pd + yellow. Podetia grayish-green, peg or rod-shaped. 
Apothecia lumpy and irregular... ... Cladonia subcariosa Nyl. On 
ground. f. squamulosa Robbins. Podetia squamulose. 

Be REE ee 0 c0 hn ada P MED E LS Cee 44 


. Podetia esorediose, grayish-green, simple or sparingly branched. KOH —, 


Pd + red....Cladonia clavulifera Vainio, f. nudicaulis Evans. On 
ground. Podetia simple or little branched, lacking squamules except 
at base. Apothecia large, one or several. 


< Podetia sorediose a. OCH LE Lou ase AE A ee ca) E 45 
i PONE TNADORD. OC EP hh i Lxx vs NR EE ERU S T 46 
. Soredia granular, or if farinose, mixed with granules................. 47 


. Tips of podetia pointed, horn-like. KOH —, Pd + red....Cladonia 


coniocraea (Flk.) Spreng. 


. Tips of podetia blunt. KOH —, Pd + red... .Cladonia fimbriata (L.) Fr. 
. Podetia covered with farinose soredia mixed with coarse granules. . Cladonia 


cylindrica Evans. 


. Soredia granular. D oe elo ke nne ERR EESE OS DEL 48 
. Podetia long, usually over 15 mm.; soredia occurring only in upper 


Daris | — vw OU EU VI oed iur Cladonia nemoxyna (Ach.) Nyl. 


48. Podetia short, less than 15 mm.; podetia sorediate throughout. . . Cladonia 
pityrea (L.) Hoffm. 
STEREOCAULON 

1. Primary thallus persistent, crustose, well developed. Stalks small, usually 
less than 1 em., simple or sparingly branched, tips densely sorediate 

Stereocaulon pileatum Ach. On rocks. 

1. Primary thallus disappearing. Stalks rather long, well developed eso- 
redigse o MGE Er DNE M ELE E 2 

2. Stalks densely covered with coralloid squamules and granules. Usually 
fruiting. Pd + orange red... . Stereocaulon dactylophyllum Flk. (Syn. 
S. coralloides Fr.) On rocks. 

2. Stalks not densely covered with coralloid squamules and granules. Rarely 
fruiting. Pd + faint sulfur yellow... .Stereocaulon evolutoides (H. 
Magn.) Frey. On rocks. Common. 

USNEA 

1. Thallus short, erect, shrub-like, sorediose, abundantly fibrillose. . . . Usnea 
hirta (L.) Wigg. On bark. 

1. Thallus short, erect, shrub-like, esorediose, less abundantly fibrillose . . Usnea 
florida (L.) Web. On bark. Fairly common. 

KEY TO THE GENERA OF ForrosE LICHENS 

1. "Thallus brightly colored, yellow or orange...........0.0.000.0 00000-0202. 2 

I TRAIGO not bnentiy coloredi vsu uso var DP RETE 

2. Thallus yellow; KOH —....Candelaria concolor (Dicks.) Arn. On bark. 
Thallus small, finely divided; lobes small, finely incised, sorediate or 
granulose. Apothecia very small, scattered, circular, yellow to orange. 
Var. effusa (Tuck.) Merrill & Burnh. Very common. Thallus reduced 
almost wholly to a powdery crust. 

2. Thallus yellowish to orange; KOH +, red to purple. .. . Xanthoria candelaria 


(L.) Kickx. Found on elm trees along roadsides. "Thallus of rather 
deeply cut, branched and imbricated lobes the margins of which are 
densely granulose or powdery, sometimes forming a thick powdery crust. 


82 HRhodor: | VoL. 60 

3. Thallus fastened by a central point below... o oiii sess 4 

3. Thallus fastened by several or more points or rhizinae................. 5 

4. Fruit bodies perithecia, immersed in thallus, globose, appearing outwardly 
as spots or small warts. Thallus pale on underside... DERMATOCARPON, 
p. 82. 

4. Fruit bodies apothecia, sessile on thallus, round or angular. Thallus 
blackish on underside...............0..0..0..0000.. UMBILICARIA, p. 84 

5. Lower surface reticulated with distinct veins....Peltigera canina (L.) 
Willd. On ground. Apothecia large, margins rolled backward, borne 
on tips of extended lobes. 

5. Lower surface not veined... unuau eie eee 6 

6. Lobes elongated, narrow. Upper cortex compact, well developed . . Anapty- 
chia hypoleuca (Muhlb.) Mass. On bark. 

6. Lobes not greatly elongated. Upper cortex loose, not compact......... 7 

7. Rhizinae usually quite numerous........ iles ese 8 

7. Rhizinae scarce or absent... liiis ee 9 

8. Spores two-celled, dark. Underside of thallus usually white, rarely brown 
or black. ee Puyscia and PyxiNE, p. 83 

8. Spores simple, uncolored. Underside of thallus most always brownish to 
blackish... nnna naaa eee PARMELIA, p. 82 

9. Thallus hollow; tips usually bursting, densely sorediate; rhizinae ab- 
sent........ Hypogymnia physodes (L.) Nyl. On bark, very common. 

9. Thallus not hollow; rhizinae searee..... llis lesus. CETRARIA, p. 82 

CETRARIA 

I. Thallus brownish... 0.000000 uli 2 

1. Thallus greenish-gray... 0.000000 0 e 3 

2. Thallus having scattered brown or black tubercles. Apothecia originating 
from under surface...........0....... Cetraria ciliaris Ach. On bark. 

2. Thallus having no tubercles. Apothecia, if present, originating from upper 
surface... . Cetraria islandica (L.) Ach. On ground in pine woods. 
Stalks strap-shaped, forking, thin, curling when dry to form a trough-like 
structure. 

3. Thallus irregularly lobed; lobes usually curled away from substrate; margins 
undulate, somewhat crenulate, covered with dense pale green soredia; 
undersurface brown. .............. Cetraria Oakesiana Tuck. On bark. 

3. Thallus lobes channeled and pitted, margins ascending, usually irregularly 


jagged and torn; undersurface pitted, white or brown near the tips, black 


at the base... eee Cetraria Tuckermanii Oakes. 
DERMATOCARPON 

Growing on rocks submerged in water.  Undersurface veined. Thallus 

lobes not pruinose when dry... . Dermatocarpon aquaticum (Weis.) Zahl. 


Found only in one location, on a rock in a stream which normally dries up 
in the summer months. 

Growing on rocks in dry situations. Undersurface not veined. Thallus 
lobes much imbrieated, pruinose when dry... . Dermatocarpon miniatum 
(L.) Mann, var. complicatum (Lightf.) Th. Fr. 


PARMELIA 


1958] Ahmadjian,—Fruticose and Foliose Lichens 83 


2. Soredia developing in rows, along cracks in the thallus or along the mar- 
fins vo s Parmelia sulcata Tayl. On bark. Very common. 
4. Boredia notan ROWS -ce ho E S LI LEAN Lo s i.d 
3. Thallus brown, with small warts breaking down into soredia..... Parmelia 
subaurifera Nyl. On bark. Very common. 
3. Thallus not brown, without warts..... 2.2.00... ee ec cc er rrr 4 
4. Thallus yellowish-green; undersurface brown or black... . Parmelia caperata 
(L.) Ach. On bark and rocks. Very common. 
4. Thallus greenish to bluish-gray; undersurface pale to drab...... Parmelia 
dubia (Wulf.) Mass. (Syn. P. Borreri Turn.) On bark. 
5. Thallus containing few blackish cilia along margins; rather broad- 
Con. ere ES Parmelia perforata (Wulf.) Ach. On bark. 
5: Phallus not elliate, not broad-lobed i. sirae i es mne 6 
6. Thallus reticulate-rimose, having a network of chinks and cracks, greenish 
TO DIADA t c eC NE. S E 7 
6. Thallus not reticulate-rimose,  yellowish-green....Parmelia conspersa 
(Ehrh.) Ach. On rocks and bark. Very common. var. stenophylla 
Ach. Lobes rather narrow and imbricated. No isidia. f. isidiata (Anzi) 
Berry. Thallus densely covered with isidia. 
7. Under surface pale. ..... Parmelia rudecta Ach. On bark. Very common. 
7. Under surface black... .Parmelia saxatilis (L.) Ach. On rocks and bark. 
Common. 
PHYSCIA and PYXINE 
ie ebhallussso0reqiose sess TEE c-r ee ELE ec 2 
1. Thallus lacking soredia, esorediose... 1.0.0.0... 0 cece eee eee eee 8 
2. Medulla yellow; apothecia lecideine. ..Pyzxine sorediata (Ach.) Fries. On 
bark. 
2. Medulla white or orange; apothecia lecanorine.......................8 
3. Lobes ascending, free from substratum... . 2.22.2 30. oe ee ee 4 
9. Lobes adnate, flattened, not ascending:......... 2.4.2. -. 2... .<.- - 5 
4. Lobes helmet or hood-shaped, with long cilia along margins; soredia within 
these hood-like pustules. KOH=..Physcia ascendens Bitt. On bark. 
4. Lobes finely incised, not hood-shaped, small, short. KOH + yel- 
low....Physcia millegrana Degel. (Syn. P. tribacia (Ach.) Nyl.) On 
bark. Very common. 
5. Lobes very thin, adglutinated to the substratum. . . . Physcia elaeina (Sm.) 
A. L. 8m. On bark. 
5. Lobes rather thick, not adglutinated to the substratum............... 6 
6. Medulla orange....Physcia orbicularis (Neck.) Potsch., f. rubropulchra 
Degel. On bark. Very common. 
a | rr rr eS 7 
7. Thallus gray or whitish, without brownish tinge... .Physcia leucoleiptes 
(Tuck.) Lettau. On bark. 
7. Thallus brownish or olive-gray....Physcia orbicularis (Neck.) Potsch. 
On bark. 
8. Thallus pruinose, having powdery appearance... .Physcia pulverulenta 
Hampe. On bark. 
8. Thallus not pruinose. Cortex KOH + yellow................ eee 9 
9. Lobes with marginal cilia........ Physcia leptalea (Ach.) DC. On bark. 
9. Lobes without marginal cilia......................... ese 10 
10. Thallus gray, with white spots. Medulla KOH + yellow....... Physcia 


aipolia (Ehrh.) Hampe. On bark. 


84 Rhodora [ VoL. 60 


10. "Thallus gray, without white spots. Medulla KOH —....Physcia stellaris 
(L.) Nyl. On bark. Very common. f. tuberculata Kernst. Lobes 
densely covered with tubercles, except at tips. var. rosulata (Ach.) Nyl. 
Lobes are wide, fan shaped at the tips. 


UMBILICARIA 

1. "Thallus pustulate, blistered or pimpled; undersurface pitted. . . . Umbilicaria 
papulosa (Ach.) Nyl. On rocks. 

1. Thallus not pustulate... 0.00.02 cee 2 

2. Undersurface having grilled or laced appearance. Thallus margins com- 
monly curling under, inrolled. Apothecia common........ Umbilicaria 
Mühlenbergii (Ach.) Tuck. On rocks. 

2. Undersurface hairy. Thallus margins not curling under. Apothecia 


rare... .Umbilicaria mammulata (Ach.) Tuck. (Syn. Gyrophora Dillenii, 
U. Dillenii). On rocks. 
XLOSSARY 


Adnate. Adhering or touching broadly to the substratum. 

A pothecium. An open cup- or saucer-shaped fruiting body in which the cavity 
is lined with a palisade-like layer of microscopic asci which bear the spores. 

Areolate. Marked out in little spaces or islands. 

Ascus. (Asci). A sac-like cell in which the spores are developed. 

Ascyphous. Podetia without cups. 

Caespitose. Growing in low tufts or patches. 

Ciliate. Having hairs or cilia. 

Coralloid. Coral-like outgrowths usually on thallus. 

Cortex. Outer layer of the thallus. 

Corticate. Covered with a continuous cortex. 

Cylindrical. Having the same diameter throughout the length. 

Dorsiventral. With two unlike sides, the upper surface differing from the 
lower surface. 

Farinose. Meal-like; mealy. 

Fibrillose. Provided with fibers. 

Flaccid. Soft and flabby; limp. 

Granulose. Covered with minute granules. Almost powdery. 

Gyrose. Convoluted; folded; wavy. 

Imbricate. Overlapping like shingles. 

Isidium.  (Isidia). A coral-like outgrowth produced on the thallus. 

KOH. Potassium hydroxide (aqueous solution—15-25%). 

Lacunose. Covered with pits or depressions. 

Lecanorine. Type of apothecium which has a rim consisting of and the same 
color as the thallus. 

Lecideine. Type of apothecium which does not have a thallus rim. 

Medulla. Made up of loosely interwoven strands or hyphae; can be observed 
when outer cortex is scraped off. 

Pd. Paraphenylenediamine (fresh saturated alcohol solution). 

Perithecium. A flask-shaped, rounded or oval fruit-body in which the spores 
are borne; lined with a palisade-like layer of asci, but with a small opening 
(ostiole) at the tip. 

Podetium. A stalk-like elevation arising from the thallus; bears the fruit- 
body. 

Primary thallus. Characterized by a horizontal and stratose thallus from 
which arises the secondary or radiate thallus termed the podetium. 


1958] Ahmadjian,—Fruticose and Foliose Lichens 85 


Pruinose. A finely powdered surface. 

Pustule. A blister or pimple-like structure. 

Pycnidium. | (Pycnidia). Small flask-shaped structures immersed in thallus. 

Reticulate. Lines, veins, or ridges crossing each other to form or give the 
appearance of a net. 

Reticulate-Rimose. A network of cracks or chinks. 

Rhizoid. A single thread-like filament (hypha) or a bundle of hyphae extend- 
ing from lower surface of the thallus and serving to attach the lichen to 
the substratum. 

Scyphus. Cup-like dilation of the podetium. 

Serrate. Saw-toothed margin. 

Sessile. With no stalk. 

Soredium. One or more algal cells surrounded or enveloped by fungal hyphae 
(threads). Masses of these when formed on the thallus give it a scattered 
or localized powdery or granular appearance. 

Spore. The reproductive cell produced by the fungus. 

Squamule. A small seale. 

Squamulose. With minute scales. 

Substrate. The substance or material to which the lichen is attached. 

Thallus. The assimilative body consisting of algal and fungal components. 

Tubercle. Any knob-like or wart-like elevation of the surface. 

Umbilicus. The central point by which means the thallus is attached to the 
substratum. 

Veined. Having raised lines or vein-like structures. 


FARLOW HERBARIUM, HARVARD UNIVERSITY 


LITERATURE CITED 


1. Arben, W. C., The Physical Features of Central Massachusetts. 
United States Geological Survey, Bulletin 760-B, 1924. 

2. ALLESCHER, ANDREAS, Rabenhorst Kryptogamen Flora, 1.7. Leipzig, 
1901-1903. 

3. Dann, Erur, On the Use of Lichen Chemistry in Lichen Systematics. 
Revue Bryologique et Lichenologique, II, 21, Paris, 1952, pp. 
119-134. 

4. ENGLER, A. and K, PnawrL, Die Natürliehen Pflanzenfamilien, 8 
hand, Lichens (Flechten), 270 pp., 127 figs, Leipzig, Engelmann, 
1926. 

5. Evans, W. A., Notes on the Cladoniae of Connecticut, IV. RHODORA, 
52: 77-123, 1950. 

6. Evans, W. A., The Cladoniae of Connecticut. Conn. Academy of 
Arts and Sciences, Vol. 30: New Haven, Conn., Yale University 
Press, June 1930, pp. 357-510. 

7. Fink, Bruce, The Lichen Flora of the United States. Ann Arbor, 
University of Michigan Press, 1935. 

8. Hane, Mason E., Lecture Notes in Lichenology, West Virginia 
University, Morgantown, 1957. 

9. Howanp, Grace E., Lichens of the State of Washington, Seattle, 
University of Washington Press, 1950. 

10. Kroc, Hitpur, Microchemical Studies on Parmelia, Saertrykk av 


86 Rhodora | Vor. 60 


Nytt Magasin for Naturvidenskapene, B. 88, Oslo, 1951, pp. 
57-85. 

11. LurrnELL, E. S., The Cladoniaceae of Virginia, Lloydia 17: 1954. 

12. Nvarina, G. G., The Lichen Book, Ridgewood, N. J., Published by 
the author, 1947. 

13. SurrH, A. L., Lichens. Cambridge, University Press, 1921. 

14. ZAHLBRUCKNER, A., Catalogus Lichenum Universalis, Vol. I-X, 
Leipzig, 1921-1940. 


THREE CHANGED AUTHORITIES FOR COMBINATIONS.—Recently, 
I had occasion to examine, page by page, the first 111 volumes of 
Curtis’s Botanical Magazine, in search of varietal names of Amer- 
ican plants for inclusion in the Gray Herbarium Card Index. In 
the course of this examination, three instances were found in which 
the current attributions of authorities and places of publication 
are incorrect. 

Aquilegia formosa, var. flavescens (S. Wats.) J. K. Henry, Fl. 
So. Brit. Columbia, 137 (1915) is antedated by Aquilegia formosa, 
var. flavescens (S. Wats.) Hook. f. in Curtis, Bot. Mag. 107: tab. 
6552 (1881). 

Mimulus luteus, var. cupreus (Regel) Hoss. in Trab. Inst. Bot. 
Farm. Buenos Aires, no. 33: 71 (1915) is antedated by Mimulus 
luteus, var. cupreus (Regel) Hook. in Curtis, Bot. Mag. 90: tab. 
5478 (1864). 

Vaccinium corymbosum, var. fuscatum (Ait.) A. Gray, Syn. Fl. 
2, pt. 1: 23 (1878) is antedated by Vaccinium corymbosum, var. 
fuscatum (Ait.) Hook. in Curtis, Bot. Mag. 62: tab. 3433 (1835).— 
ROBERT C. FosrkEn, GRAY HERBARIUM. 


SOME IDENTITIES IN HALESIA (STYRACACEAE) 
R. K. GODFREY! 


In western Florida, from the Tallahassee Red Hills area west- 
ward, the two-wing silverbell, Halesia diptera Ellis, comprises two 
populations: a larger flowered population which in general grows 
in mixed woodlands of upland slopes in the Tallahassee Red Hills, 
mixed woodlands of bluffs and ravine slopes along the Apalachi- 

1 Expenses incurred in field work contributing to this research were met with funds made 
available to the author by a grant from the National Science Foundation (G-2010). Her- 


barium assistance was made possible by a grant from the Research Council, Florida State 
University. 


Rhodora Plate 1231 


Pecks Prans Vol tg TAB MB oe 


A HALESIA i wevapiine fert po E 
B HALKSIA / Apron fom tm n 


Halesia tetraptera Ellis, fig. A.  Halesia diptera Ellis, fig. B. After Ellis. 


1958] Godfrey,—Identities in Halesia (Styracaceae) 87 


cola and Chipola Rivers, an | mixed woodlands of the uplands hav- 
ing limestone outerops in the vicinity of Marianna; a smaller flow- 
ered population which grows in the broad floodplain forests of the 
Choctawhatchee and Escambia Rivers and in lesser floodplains of 
smaller streams between. I have, as yet, no reason to believe that 
the larger flowered form is of more widespread distribution than 
from the Tallahassee to the Marianna area. Neither have I, to 
date, encountered any of the smaller flowered form within this 
limited range. The smaller flowered one, in western Florida oc- 
curring from about the Choctawhatchee River westward, is a 
part of a much more extensive population extending westward to 
Texas and northeastward to the Savannah River (or thereabouts). 

Aside from habitat and distribution differences, the only 
morphological differences between these two-wing silverbells is 
that of flower size. This is conspicuous as one views the trees, 
those of the larger flowered form being nearly twice the size (the 
form and proportions being the same) of the smaller flowered one. 
However, when flowering material is carefully pressed, the size 
difference is less strikingly apparent in herbarium specimens. The 
flowers shrink considerably in drying. 

Although in the non-flowering condition, the two-wing silverbells 
are not distinguishable, it seems advisable to give them varietal 
designations. This necessitates, of course, ascertaining what Ellis 
may have had when he described Halesia diptera. The species was 
described by Ellis in the Philosophical Transactions of the Royal 
Society, London, 51: 931, t. 22, fig. B (1791). This is in the paper 
in which he also described (and figured much more fully) H. 
tetraptera the identity of which is considered below. At the con- 
clusion of his discussion of H. tetraptera, Ellis wrote (of H. diptera) 
as follows: 

“About two years ago, I received from Governor Ellis of Georgia another 
species of this tree, which was sent him by Mr. De Brahme, from Augusta 
in Georgia, three hundred miles up the river Savannah. 

“The fruit of this kind has two wings, as described in the plate, at B." 

'There follows a diagnosis of Halesia, then the following species 
designations: 

The species are, 


TETRAPTERA. 1. HALEsIA fructibus 
membranaceo-quadrangulatis. 
DIPTERA. 2. Harresia fructibus alatis. 


88 Rhodora [ VoL. 60 


From this, and in the light of my present knowledge of the dis- 
tribution of the smaller and larger flowered forms of Halesta dip- 
lera, it may be inferred that the species is based on material of the 
smaller flowered one which thus becomes Halesia diptera Ellis var. 
diptera. Fresh flowers of this are 1.0-1.5 cm. long at anthesis. 
The width of the corolla varies because the petals spread vari- 
ously. The individual petals are oval, oblong, or obovate, 1.0- 
1.5 em. long and 0.8-1.0 cm. broad at their broadest places, the 
tips obtuse or rounded. The androecium length is equal to that of 
the petals, the filaments united at base. 

The larger flowered form I designate as Halesia diptera var. 
magniflora, var. nov. Var. diptera similis sed floribus maioribus. 
Floribus 2-3 em. longis, 1.0-1.5 cm. latis. Petalis 2-3 em. longis, 
1.0-1.5 cm. latis, ovalibus, oblongis, ad obovatis. Type specimen: 
deciduous woods, ravine slope, 1.5 miles east of Tallahassee, 
Florida, Godfrey no. 54434 (Herbarium of Florida State 
University). 

Halesia tetraptera Ellis (1. e. 030-931, t. 22, fig. A) was described 
from material sent to England from along the banks of the Santee 
River in South Carolina. The plate (fig. A) accompanying the 
description, which is here reproduced, admirably portrays flower- 
ing material and a fruiting branch. ‘This seems to me clearly and 
unequivocally identifiable with the little silverbell of the coastal 
plain, Halestia parviflora Michaux (Flora Boreali-Americana 2: 
40. 1803), and not with the much larger flowered silverbell of the 
uplands and interior, Halesta carolina L., with which it has long 
been identified. 

Halesia tetraptera Ellis is the earlier name and the one which 
should be applied to the tree which we are currently calling H. 
parviflora — Michx.—DbEPARTMENT OF BIOLOGICAL SCIENCES, 
FLORIDA STATE UNIVERSITY, TALLAHASSEE, FLORIDA, 


Volume 60, no. 710, including pages 33-60, was issued 19 March, 1958, 


: TT 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR, 

IVAN MACKENZIE LAMB 


Vol. 60 April, 1958 No. 712 
CONTENTS: 
Determination of Polyploidy from Herbarium Specimens. 
Robert P. Celarter-and K. D. Meha... casn is wc cs ec sca 89 
The Name Lepanthes Turialvae: A Source of Confusion. 
Richard Evans Schutes: sos see a ss pee ee ss cs 97 
Chromosome Numbers in the Genus Krameria: Evidence for 
Familial Status. B. L. Turner. cc ee es occ acs 101 
A Taxonomic Study of the Genus Physalis in North America North 
of Mexico. U. T. Waterjal so tae ceo es nt ce ee 106 
New Illinois Carex Records. Egbert W. Fell................... 115 


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JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 April, 1958 No. 712 


DETERMINATION OF POLYPLOIDY FROM 
HERBARIUM SPECIMENS 


ROBERT P. CELARIER AND K. L. MEHRA 


In the evolutionary history of the flowering plants there are 
several biological phenomena known to be of major consequence. 
Foremost among these, and perhaps the best understood, is 
polyploidy. A brief survey of any of the recent compilations of 
chromosome numbers of the Angiosperms (Löve and Löve, 1948, 
Delay, 1951; Darlington and Wylie, 1955) will suffice to show the 
frequency of this phenomenon. There are literally hundreds of 
examples of so called intraspecific polyploids reported, not to 
mention the even more frequent condition of interspecific 
polyploidy. 

The importance of polyploidy in the critical evolutionary fields 
of taxonomy, geobotany, hybridization, mode of reproduction, 
etc. has been shown and discussed by many workers (Löve, 1951; 
Löve and Löve, 1949; Gustafsson, 1947; 1948; Muntzing, 1936; 
Stebbins, 1940; 1950; Darlington, 1956, etc.). Not only is the 
importance of polyploidy well known but much is understood 
concerning its biological mechanisms of operation. 

Because of the obvious significance of polyploidy in both con- 
tinuous and discontinuous variation of plants, it is a factor that 
cannot easily be dismissed in any detailed study involving the 
relationships of species or their modes of origin. 

Determination of chromosome numbers is, however, a time 
consuming operation, and becomes virtually impossible for 
monographers who deal principally with non-living herbarium 
materials. Some effort has been made to overcome this handicap 
and recently Khoshoo (1955) has been able to study chromosomes 


90 Rhodora [Vor. 60 


from herbarium material in Impatiens. However the techniques 
are rather laborious, the results far from the best, and the extent 
to which the technique is applicable is not yet known. 

Numerous studies have been conducted that attempt to cor- 
relate morphological conditions with degree of ploidy (see Steb- 
bins, 1950 for review), and some workers believe that there is 
almost always some correlation (Lóve, 1951). Although the 
general conclusion from these studies is that there are no univer- 
sal criteria, nevertheless there are certain characters that have 
rather general application. Foremost among these is cell size. 

From herbarium material there are usually easily available two 
types of cells (pollen grains and guard cells of the stomata). 
Since there is often some overlapping between cell size and degree 
of ploidy, it is desirable, and sometimes essential, to study the 
cell size of both before drawing conclusions. 

In general, pollen grains are easily studied, but the conven- 
tional method for studying guard cells requires a pretreatment of 
the leaf, followed by scraping or stripping (deWet, 1954). In any 
event some portion of the specimen is mutilated or destroyed, a 
condition that is very undesirable especially with valuable speci- 
mens such as types. 

In this report an impression method is described for the study 
of the guard cells of the stomata that is quick, reliable, and causes 
no damage to the specimen. This method is based on the prin- 
ciples of impression long used in paleontology, and with the use 
of some of the modern plastics gives very desirable results. 
Somewhat similar methods have been used by plant pathologists 
(Long & Clements, 1934; Husain, 1956) to detect the open or 
closed condition of the stomata. 

The procedure is simply to mix cellulose nitrate! in acetone 
until a viscous solution is obtained. When the constancy of 
the solution is such that it spreads smoothly with a camels hair 
brush, paint the surface of the leaf to be studied with the solution. 
This will work better if the leaf surface has been previously 
cleaned with acetone. The solution is allowed to dry thoroughly 
and is then peeled off with tweezers. This plastic strip is then 


! Jacobo Ortega Castro of this department, in a study involving the relationship between the 
opening and closing of the stomata in wheat and leaf rust infection, has used cellulose acetate 
and collodion with equal success. His work shows that at different temperatures different 
plastics are preferable. 


1958] Celarier and Mehra,—Determination of Polyploidy 91 


floated in a drop of water on a slide, covered with a cover slip, 
and is ready for study under the microscope. 

Cell impressions prepared by this method are usually distinct 
(figs. 1-8) and measurements can be made with considerable 
confidence. 

RESULTS 

In the present study several grass species complexes of the tribe 
Andropogoneae were analyzed. A rather detailed study was made 
of the Dichanthium annulatum complex, which included diploids 
(2n = 20), tetraploids (2n = 40), and hexaploids (2m = 60). 
Also studied, but in less detail, were tetraploids, pentaploids, and 
hexaploids of the Bothriochloa ischaemum complex, and tetra- 
ploids and hexaploids of the B. zntermedia and B. pertusa com- 
plexes. "The chromosome numbers of all accessions used in this 
study were previously determined (Celarier, 1957; Celarier and 
Harlan, 1955; Celarier, Mehra, and Wulf, in press, and 
unpublished). 

Pollen grains and stomata guard cells were studied from both 
fresh material and herbarium specimens and the results are given 
in tables 1 and 2. Although most specimens were only three or 
four years old, it seems likely that, under proper storage condi- 
tions, only a negligible amount of change would be expected with 
the age of the specimen. 


D. ANNULATUM COMPLEX 


In the present report three diploid, eight tetraploid, and two hexaploid 
accessions were studied. The data are presented in table 1, and figures 1-8 
show their general appearance. 

Pollen grain size was quite variable in all accessions with a range of 
approximately 10u. However the means were similar in all accessions of 
one ploidy level, and quite different between polyploids (figs. 1-3). The 
diploids means ranged from 32.0 to 33.0u, the tetraploids from 36.2 to 
39.9u, and the hexaploids from 42.9 to 48.7u. Pollen grains from herbarium 
specimens were almost always smaller than fresh material but usually the 
mean values were of less than one micron difference, 

There was also variation in stomata guard cell size but it was much less 
than in pollen grains and was in general, less than five microns. Again the 
mean values were quite distinct at the different ploidy levels. In the fresh 
material the means in the diploids varied from 23.7 to 24.8u, in the tetra- 
ploids from 30.0 to 32.0u, and the hexaploids varied from 36.8 to 45.9. 
The same kind of variation was seen in the herbarium specimens but guard 
cells were in all cases considerably s maller than in fresh material (figs. 4-6). 


[Vor. 60 


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Celarier and Mehra,—Determination of Polyploidy 


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94 Rhodora [Vor. 60 


Although the diploids, tetraploids, and hexaploids could be easily dis- 
tinguished from one another in the herbarium materials it is obvious that 
comparisons between herbarium and fresh materials cannot be made until 
a correction factor is established. 


B. ISCHAEMUM COMPLEX 

Two accessions each were studied in the tetraploids, pentaploids and 
hexaploids (table 2). As in D. annulatum there is considerable variation in 
pollen grain size (ca. 10-154) whereas the range in variation in the guard 
cells is small (less than 54). The means however were rather constant for 
both, but different in different ploidy levels. 

In the tetraploids the means of pollen grain size ranged from 36.7 to 
37.1u for fresh materials and 35.4 to 36.64 for specimens. The pentaploids 
ranged from 37.8 to 38.2u for fresh materials and 35.5 to 39.3u for speci- 
mens, and in the hexaploids the range was 41.9 to 43.9u for fresh material 
and 40.9 to 43.9u from specimens. 

Stomata guard cells were also distinct but as in D. annulatum showed a 
big difference from fresh material to specimens. 

In the tetraploids the means ranged from 25.0 to 28.74 in fresh materials 
and 18.0 to 19.04 in specimens. The pentaploids ranged from 29.0 to 
30.2u in fresh materials and 20.6 to 21.3u in specimens, and the hexaploids 
ranged from 30.7 to 31.8u in fresh material and 21.8 to 22.04 in specimens. 

In B. ischaemum it seems that pentaploids cannot be easily separated 
from hexaploids on guard cell size alone but by the use of both pollen grain 
and stomata guard cells the separation is fairly reliable. 


B. INTERMEDIA COMPLEX 


In this species only two tetraploids and two hexaploids were used. The 
tetraploids were readily distinguishable from the hexaploids with both 
pollen grain size and stomata guard cell size. "Variation in both was rather 
similar to that seen in D. annulatum and B. ischaemum. 

The tetraploids had a range in mean pollen grain size of 34.7 to 36.9u 
in fresh materials and 34.4 to 35.0u in specimens; whereas the hexaploids 
ranged from 39.7 to 42.0u in fresh material and 39.7 to 41.9u in specimens. 

The range of the means in guard cell size were 25.5 to 25.8u in fresh 
materials and 14.7 to 15.2u in specimens for the tetraploids and for the 
hexaploids were from 29.1 to 32.8u for fresh materials and 19.6 to 20.1u 
for specimens. 

B. PERTUSA COMPLEX 
In this species complex two tetraploids and two hexaploids were used 


and the results were similar to those found in the other species. 
The range in pollen grain means was 35.8 to 37.6u for fresh material 


Figs. 1-8. Pollen grains and stomata guard cells in Dichanthium annulatum. Figs. 1-3. 
Pollen of the three ploidy levels. X300. Fig. 1 diploid. Fig. 2 tetraploid. Fig. 3 hexaploid. 
Figs. 4-6. Comparison of stomata guard cells from fresh mounts and plastic peels of herbarium 
specimens in the three ploidy levels. X1350. Fig. 4 diploid. a. fresh material. b. plastic 
peel of specimen. Fig. 5 tetraploid. a. fresh material. b. plastic peel of specimen. Fig. 6. 
hexaploid. a. fresh material. b. plastic peel of specimen. Figs. 7-8. Comparison of plastic 
peel of fresh material (Vig. 7) with plastic peel of specimen (Fig. 8) in the hexaploid. X300. 


Rhodora Plate 1232 


Fig. 1-8. For explanation see opposite page. 


1958] | Celarier and Mehra,—Determination of Polyploidy 95 


and 35.0 to 36.5u for specimens in the tetraploids, and 39.4 to 44.34 for 
fresh material and 37.1 to 42.2u for specimens in the hexaploids. 

The stomata guard cell size was also distinct with means ranging from 
24.3 to 25.8u in fresh material and 13.9 to 14.24 in specimens for the tetra- 
ploids; whereas, the hexaploids ranged from 30.0 to 34.9u in fresh material 
and 20.2 to 21.64 in specimens. 


DISCUSSION AND CONCLUSIONS 

It has been shown that both pollen grain and stomata guard cell 
size are fairly reliable indicators of the degree of ploidy in several 
of the Old World species of the genera Bothriochloa and 
Dichanthium. It has also been shown by Gould (1957) that 
pollen grain size is useful in determining the degree of ploidy in 
several of the American species of Bothriochloa. 

In the species studied there was no difficulty in distinguishing 
between diploids, tetraploids and hexaploids. However in B. 
ischaemum the differences between the pentaploids and hexaploids 
were not so distinct, but by the use of both pollen grain and guard 
cell sizes a fairly reliable conclusion could be drawn. 

In general it was possible to place the materials studied in their 
proper ploidy levels by pollen grain and guard cell size regardless 
of the species involved. However there were exceptions to this, 
such as the tetraploid D. annulatum A-4099 with pollen grains 
39.9u and guard cells of 32.04 and the hexaploid B. intermedia 
A-4597 with pollen grains of 39.74 and guard cells of 32.8. 

The impression technique for measuring guard cells is shown 
to be quite reliable but the actual measurements were in all cases 
much less than those made from fresh material. In order to 
determine what portion of this decrease in length was due to the 
technique and what portion was due to the drying of the speci- 
mens, measurements were also made from plastie strips taken 
from fresh material. 

In the diploid D. annulatum A-3242 the mean guard cell meas- 
urements from fresh material was 24.34 whereas the plastic 
strip measurements from specimens was 13.64. Plastic strip 
measurements of fresh material of this accession were found to 
be 20.44. From this it is seen that a considerable portion of this 
decrease in size is due to the technique itself but that most of it is 
probably due to the shrinkage in drying of the specimens. A 
similar condition was found in the hexaploid (figs. 7-8) but only a 
few measurements were made. 


96 Rhodor: [Vor. 60 


These studies seem to warrant certain recommendations in the 
procedures used in studies where it is desirable to determine 
chromosome numbers from herbarium specimens. The following 
appear to be significant: 

1. Pollen grain and stomata guard cell size are usually reliable 
indicators of polyploidy, and the use of both would be expected 
to give much more dependable results than either alone. 

2. Actual chromosome counts should be made from at least a 
few plants of several polyploid levels. Pollen grain and guard 
cell measurements from these plants can serve as a standard. 

3. Data should be calculated in terms of ranges and means. 
This seems to be especially important in studies of pollen grains. 

4. Guard cells from herbarium specimens can be reliably meas- 
ured by the impression technique but when compared with fresh 
material a correction factor must be taken into account to offset 
the shrinkage. 

5. Conclusions regarding chromosome number based on cell 
size should be transferred to a second species with extreme cau- 
tion, unless some chromosome counts of the second species have 
been made so that a standard can be established. 

6. Data concerning cell size would be a valuable addition to a 
monograph even if the chromosome numbers of the taxa involved 
are not known, in that they may offer a suggestion of polyploidy 
and will be available if cytological studies are made in the future. 


SUMMARY 


Data are presented that demonstrate a correlation between the 
degree of polyploidy and size of pollen grains and stomata guard 
cells in four species complexes of the grass genera Dichanthium 
and Bothriochloa. These studies were made both from living 
material and dried herbarium specimens. 

An impression technique using plastic strips is outlined for the 
study of stomata guard cells from herbarium specimens. This 
technique gives reliable measurements without damage to the 
specimens. 

Some of the limitations to the use of cell size as a gauge of poly- 
ploidy are discussed and certain recommendations are offered 
based on present studies. DEPARTMENT OF BOTANY AND PLANT 
PATHOLOGY, AGRIC. EXP. STA., OKLAHOMA STATE UNIVERSITY, 
STILLWATER. 


1958] Schultes,—Lepanthes Turialvae 97 


LITERATURE CITED 


CüLARIER, R. P. 1957. The Cyto-geography of the Bothriochloa ischaemum 
complex. II. Chromosome behavior. Amer. Jour. Bot. 44: 729-738. 
-———-—- AND J. R. Hartan. 1955. Studies on Old World Bluestems. 
Okla. Agr. Exp. Sta. Tech. Bull. T-58: 1-31. 
- ———— —-, Menna, K. L. AND M. L. Wurr. (press). Cytogeography of the . 
Dichanthium annulatum complex. Brittonia. 10: 

DinuiNaGTON, C. D. 1956. Chromosome Botany. London, England. 

—-—-——— —- AND A. P. Wye. 1955. Chromosome Atlas of Flowering 
Plants. London, England. 

DkrAv, C. 1951. Nombres Chromosomiques chez les Phanérogames. Rev. 
de Cyt. et biol. végét. vol. 12. 

DEWET, J. M. J. 1954. Stomatal size as a cytological criteria in Danthonia. 
Cytologia 19: 176-181. 

Gourp, F. W. 1957. Pollen size as related to polyploidy and speciation in 
the Andropogon saccharoides—A. barbinodis complex. Brittonia 9: 71-75. 

GUSTAFSSON, i 1947. Apomixis in higher plants. III. Biotype and species 
formation. Lunds Univ. Arsskr. 44: 183-370. 

—————————. 1948. Polyploidy, life-form, and vegetative reproduction. 
Hereditas 34: 1-22. 

Husain, S. M. 1956. Studies of competitive ability of certain races of 
wheat leaf rust. M.S. Thesis. Oklahoma A & M College. 

Kuosnoo, T. N. 1956. Chromosomes from herbarium sheets in /mpatiens. 
Stain Tech. 31: 31-33. 

Lona, F. L. AND F. E. CLEMENTs. 1934. The method of collodion films for 
stomata. Amer. Jour. Bot. 21: 7-17. 

LóvE, A. 1951. Taxonomical evaluation of polyploids. Caryologia 3: 
263-284. 

———— ———- AND D. Lóvk. 1948. Chromosome numbers of Northern plant 
species. Icel. Univ. Inst. Appl. Sci., Dept. Agric. Rep. B. 3: 1-131. 

— AND ———— —————. 1949. The geobotanical significance of 
polyploidy. I. Polyploidy and latitude, Portug. Acta Biol. (A) R. B. 
Goldschmidt vol. 273-352. 

Muntanc, A. 1936. The evolutionary significance of autopolyploidy. 
Hereditas 21: 263-378. 

STEBBINS, G. L. 1940. The significance of polyploidy in plant evolution. 
Amer. Nat. 74: 54—66. 

1950. Variation and evolution in plants. New York. U.S.A. 


THE NAME LEPANTHES TURIALVAE: A SOURCE OF 
CONFUSION 
RICHARD EVANS SCHULTES 


One of the most puzzling concepts in the orchid genus Lepanthes 
has been L. turialvae Rchb. fil., described from material collected 
in Costa Rica. The binomial has been applied to a bewildering 
array of species which obviously represent different concepts and 
has, as a result, become a rather convenient, albeit an inexact, 


98 Rhodora [ Vor. 60 


receptacle to which are referred many Middle American collections 
the identification of which might pose difficulties or doubts. 

During my investigation of the species of Lepanthes known from 
Mexico, the name Lepanthes turialvae, with its present vague 
interpretation, continually confounded the study. Critical exam- 
ination of what has been called Lepanthes turialvae in the literature 
and herbaria convinces me that, whichever of the several concepts 
is followed, it is not represented in Mexico. Nevertheless, it would 
seem advisable to present the following notes which have grown 
out of my study of Lepanthes turialvae, so that they may be made 
available to future workers in the genus. 


Lepanthes turialvae Reichenbach fil. in Bonpl. 3 (1855) 225. 

Original description :“ Lepanthes Turialvae ( Effusae) : similis L.cochleari- 
formi Sw. vaginis arctis, ostio tantum angusto microscopice muriculatis, 
folio ovato acuto bene limboso, racemi pectinati bracteis parvis muricatis, 
sepalo inferiori alte bifido, tepalis incisione triangula extrorsa acuta 
bilobis, labelli lobis obtuse triangulis. Stengel drei bis vier Zoll hoch 
mit Einschluss des zollangen Blatts. Bluthe nach Hrn. Dr. Oersted's 
zeichnung beschreiben. Sepalen dreieckig fleischroth. Tepala gelblich, 
obserseits mit Purpurroth auf Innenecke. Lippenlappen und Saule 
purpurroth. Turialva in Costarica, 3000'. Oersted.” 

In the same year, Reichenbach (Xen. Orch. 1 (1855) 151, 156, 
t. 50, fig. V, 15-16) published a second (Latin) and a third 
(German) description, differently worded but referring essentially 
to the same concept. In connection with these descriptions, he 
published drawings showing (V) a leaf (with its sheaths) and in- 
florescence (with one flower), (15) a flower and (16) a bract. 
Heichenbach intimated (loc. cit. 151) that the description was 
based upon Oersted's collection in the Copenhagen Herbarium 
(“Vid. sp. sice. in herbario Hafniensi’’). 

Through the kindness of Dr. O. Hagerup, director of this her- 
barium, we were able to borrow what is probably the type of 
Lepanthes turialvae and a water-color of the flower made by Oer- 
sted himself from the living plant in the field. The binomial 
"Lepanthes Turialvae Rehb.” is written under the floral drawing 
in Reichenbach's own hand. There is no flower present on the 
specimen. Probably none was ever available to Reichenbach, for 
he stated clearly (Bonpl. 225) that his description of the flower was 
based on Oersted’s drawing. Certainly, Reichenbach’s rather in- 
adequate sketches of the flower match very closely the Oersted 
water-color. 


1958] Sehultes,—Lepanthes Turialvae 99 


We have also, thanks to the cooperation of Dr. K. H. Rechinger, 
borrowed for study from the Reichenbach Herbarium in Vienna 
a sheet (No. 54553) on which there is a sterile specimen and a 
colored sketch of the flower. This sheet from Vienna is likewise 
labelled “Lepanthes Turialvae Rehb." 

The sterile collection from Vienna is said to represent Oersted's 
collection made in Turrialba, Costa Rica, at 3,000 feet altitude. 
It is here, however, that confusion enters the pieture. Neither the 
two detached leaves nor the sheaths covering the secondary stem 
represent, in my opinion, the same species as that which we have 
thought to be the type. But the colored sketch of the flower is 
identical with that in the Copenhagen material and, accordingly, 
corresponds to the floral diagram published by Reichenbach. 

Reichenbach's original description of the sterile parts seems not 
to have been based on the suborbicular-ovate and apically tri- 
dentate leaf which is found in Copenhagen, but rather on the el- 
liptie and very acute leaves attached to sheet No. 54553 from 
Vienna. His second description (Xen. Orch. 151) definitely refers 
to the Copenhagen material, for he stated that the leaf was “apice 
attenuato brevissimo tridentatum.” 

There is in the Reichenbach Herbarium a sheet (No. 54553) 
which Reichenbach himself labelled “Lepanthes Turialvae.” The 
collection was made in Turrialba by Wendland: Wendland 549. 
This sheet has a habit sketch of the plant and a very careful draw- 
ing of the flower. Mr. Elmer W. Smith recently prepared a larger 
drawing from a flower taken from this collection, boiled and floated 
out in water; his drawing agrees in all essentials with Reichenbach’s. 
In Wendland 549, the anterior lobe of the petals is triangular, the 
posterior obliquely subquadrate; whereas the petals shown in the 
type-drawing of the flower of Lepanthes turialvae have both lobes 
elliptic-lanceolate. The petal-lobes in the former are not spread- 
ing and, consequently, have no sinus between them; those of the 
latter are shown as very spreading with a conspicuous sinus. 

Reichenbach not only annotated Wendland 549 as Lepanthes 
turiclvae. In an article treating of the orchids collected by Oersted, 
he (Beiträge zu einer Orchideenkunde Central Americas (1866) 
57, 90, t. 10, figs. III, 16) illustrated as Lepanthes turialvae what is 
obviously the orchid represented in the colleetion Wendland 549. 
This illustration is wholly distinct from that published in Xenia 
Orchidacea eleven years earlier. 


100 Rhodora [Vor. 60 


In view of 1) the apparent diserepancy in sterile parts between 
the Copenhagen and the Vienna material collected by Oersted, 
both of which Reichenbach considered to represent Lepanthes 
turialvae and 2) his identification of Wendland 549 as L. turialvae, 
we must agree that Reichenbach himself had either an extremely 
vague or else a confused conception of the morphological characters 
on which L. turialvae was based. 

An examination of the Middle American material labelled 
Lepanthes turialvae in the Orchid Herbarium of Oakes Ames indi- 
‘ates the extent of the confusion attendant upon the use of this 
binomial, for there are clearly several concepts involved which 
only a monographie study of the genus can clarify. Even in modern 
taxonomic and floristic treatments, one can sense this confusion 
and doubt. Schweinfurth (in Standley “Flora of Costa Rica" in 
Field Mus. Nat. Hist. Bot. Publ. 391 (1937) 245) followed what he 
thought to be Reichenbach’s concept of the species as shown by his 
citation as basic material both the Oersted and the Wendland col- 
lections. Pertinent to this point, perhaps, is Schweinfurth’s ob- 
servation (loe. cit. 242) that he considered Lepanthes Brenesii 
Schltr. to be “very close to, if not the same as, L. turialvae." In 
1946, L. O. Williams (in Woodson & Schery “Flora of Panama" in 
Ann. Mo. Bot. Gard. 33 (1946) 84) stated that “Lepanthes turial- 
vae, as now delimited, is possibly an aggregate of several species.”’ 
Extreme confusion marks the treatment of Lepanthes turialvae by 
Ames & Correll (“Orchids of Guatemala" in Fieldiana, Bot. 26 
(1952) 196, 204). Their description is so very broad that it would 
embrace a number of species. Furthermore, there is a discrepancy 
between the characters used to key out Lepanthes turialvae (e.g. 
“Lip without a ciliate apieule or pubescent midlobe . . .") and the 
detailed description of the concept (e.g. lip “with a minute pu- 
berulent apicule in the sinus"). 

In view of this extreme confusion which, as we have seen, goes 
as far back as the original description, I suggest that the binomial 
Lepanthes turialvae Rchb. fil. be rejected as a nomen confusum in 
accordance with Articles 63 and 65 of the /nternational Code of 


1 There is here a discrepancy between the French and the English versions. The French 
version (as well as the Spanish) uses the word ‘‘confusion’’ where the English (and the German: 
"Irrtum") employs "error": “Un nom est a réjéter . . . s'il est une source de confusion." Lan- 
jouw has stated in the preface of the International Code . . . that “it was decided [by the Nomen- 
clature Section] that, should there be any inconsistency between the versions, the English 
one would be regarded arbitrarily as correct.” 


1958] Turner,—Chromosome Numbers in Genus Krameria — 101 


Botanical Nomenclature as adopted at Paris in 1954: (Article 63) 
“A name must be rejected . . . if it is a source of error"! and (Article 
65) “A name must be rejected if it is used in different senses and so 
has become a long-persistent source of error." It seems to me that 
no purpose would be served by trying to retain a name which 
apparently can never be clarified with satisfaction. However, in 
view of the fact that monographie studies in Lepanthes will shortly 
be initiated, I merely offer this as a suggestion and leave formal 
action to the monographer. 


CHROMOSOME NUMBERS IN THE GENUS KRAMERIA: 
EVIDENCE FOR FAMILIAL STATUS 


B. L. TURNER 


The genus Krameria is composed of about 20 species of peren- 
nial herbs and shrubs, most of which occupy the warmer desert 
or semi-desert regions of North and South America (Britton, 
1930). Since its initial description in 1762 the genus has been a 
taxonomic ‘“‘problem”, both as to rank and phyletie position. 
Some workers have recognized it as the single genus of the family 
Krameriaceae (Chodat, 1890; Small, 1903; Britton, 1930; Abrams, 
1944; Cronquist, 1957; ete.); other workers have assigned the 
genus subfamilial rank within the Leguminosae (Benson and 
Darrow, 1954; Benson, 1957); while still others have relegated the 
group to merely tribal status within the subfamily Caesal- 
pinioideae of the Leguminosae (Taubert, 1894; Capitaine, 1912; 
ete.). 

Such differing taxonomic treatments are not particularly dis- 
turbing since most of the workers mentioned above are more or 
less in agreement that the relationship of Krameria lies within or 
close to the Leguminosae and in particular to the tribe Caesal- 
pinioideae. However, there are serious doubts as to its phyletic 
position. It is interesting to note that while such an eminent 
worker as Taubert in Engler and Prantl’s PFLANZENFAMILIEN 
treats the genus as a tribe within the subfamily Caesalpinioideae 
of the order Rosales, Hutchinson (1926) places the genus in the 
family Polygalaceae of the order Polygalales, quite removed from 
the Leguminosae proper. Indeed, Taubert had enough confidence 


102 Rhodora [ Vor. 60 


in his treatment to relate Krameria specifically to the tribes 
Cassieae and Eucaesalpinieae and so placed Krameria as tribe 6 
between these two taxa. Hutchinson does not give reasons for 
the inclusion of Krameria in the Polygalaceae, but precedence for 
such a treatment may be found in Bentham and Hooker (1862) 
who also placed it in the Polygalaceae. "The latter authors, in 
treating this family, listed Krameria last among a group of 
“venera affinis aut exclusa, v. dubia." Hallier (1912) also viewed 
the relationship of Krameria as being with the Polygalaceae but 
assigned it familial status. 

Kunz (1913) has given the most detailed study of the problem 
to date. After a review of the literature and as a result of his 
own observations on exomorphie and anatomical characteristics 
he concluded that Krameria did not belong within the Caesal- 
pinioideae but rather should be treated as a distinct family. He 
did not attempt to show phyletic position, but he did indicate 
that Krameria was perhaps closer to the Leguminosae than the 
Polygalaceae. 

Since Kunz's excellent study only a few published facts have 
been added. Heimsch (1942), using anatomical criteria, con- 
sidered the position of Krameria with respect to the Leguminosae 
and Polygalaceae. He concluded that Krameria, on the basis of 
wood structure, was closer to the latter family. However, Erdt- 
man (1944), on the basis of pollen morphology, briefly commented 
on the unnatural position of Krameria when placed in the Poly- 
galaceae and stated that it belonged to the Caesalpinioideae of the 
Leguminosae. Dr. John Dwyer (personal communication), after 
a broad study of floral types within the Caesalpinioideae, has 
concluded that Krameria does not belong within this subfamily, 
though he has no set opinion of its phyletie position. 

In the present paper chromosome evidence has been used to 
evaluate the position of Krameria with respect to the Caesal- 
pinioideae. Unfortunately, chromosomal information is not ade- 
quate to permit comparisons with the supposed extra-leguminous 
relatives of Krameria, so little can be added to the controversy 
regarding phyletie position. 

K. grayi and K. ramosissima are small shrubs of semi-desert 
and desert regions of North America, while K. lanceolata is a 
widespread, common perennial herb which occurs throughout the 


1958]  Turner,—Chromosome Numbers in Genus Krameria 103 


CHROMOSOME NUMBERS 


Meiotic chromosome counts! were obtained for three species of Krameria 
as enumerated below: 


Species Source n 
Krameria grayi TEXAS, Terrell Co.: 10 mi. east Sanderson. 6 
Rose & Painter B.L.T. 3927 
Krameria lanceolata TEXAS, Coryell Co.: 3 mi. north Cooperas 6 
Torr. Cove. B.L.T. 3811 

^ 4: TEXAS, Kinney Co.: 12 mi. northeast Bracket- 6 
ville. B.L.T. 3803 
i: p TEXAS, Leon Co.: Marquez Dome. M. C. 6 
Johnston et al. 54994 
fe $ TEXAS, Val Verde Co.: 10 mi. northwest 6 
Langtry. B.L.T. 3771 
Krameria ramosissima TEXAS, Kinney Co.: 10 mi. southeast Bracket- 6 
(Gray) Wats. ville. B.L.T. 3874 


Southwestern United States and Mexico. All collections 
examined proved to be diploid with n = 6. Since the chromo- 
somes in the first division of meiosis are exceptionally large, 
showing three or more chiasmata at metaphase (fig. 1), two- 
dimensional camera lucida drawings and photography become 
diffieult.? Best counts are obtained from Division II of meiosis, 
when the chromosomes are less massive and thus flatten more 
easily. As indicated in figures 2 and 4, these chromosomes have 
nearly medium centromeres. 


DISCUSSION 


Taubert in Engler and Prantl’s PFLANZENFAMILIEN treated 
Krameria as the sole genus of the tribe Kramerieae, placing it after 
the tribe Cassieae of the Caesalpinioideae. By reference to floral 
morphology, Taubert (footnote, p. 166) explicitly reckoned its 
relationship to be with this latter tribe. 

Since the inclusion of Krameria within the Leguminosae should 
depend upon the total similarities it shares with members of the 
Caesalpinioideae, it seems appropriate to examine the chromoso- 
mal evidence bearing on this presumed relationship. As indi- 


1 Buds were killed and fixed in a mixture of 4 chloroform: 3 absolute alcohol: 1 glacial acetic 
acid. Anthers were squashed in acetocarmine 3-14 days after collection. Voucher sree 
are deposited in The University of Texas Herbarium, Austin, Texas. 

2 The meiotic chromosomes of Krameria rank among the largest known within the nda 
Covas and Schnack (1946) and Baldwin and Speese (1957) have documented somewhat larger 
meiotic chromosomes for two parasitic species of the Loranthaceae (Psittacanthus cuneifolius 
and Phoradendron flavescens). Except possibly for those of the well known species of Paeonia, 
these species have the largest meiotic chromosomes of any dicot known to the present writer. 


104 Rhodora [Vor. 60 


cated above, Krameria is, so far as known, unibasic with a num- 
ber of x = 6, its meiotic chromosomes being especially noteworthy 
for their very large size and several chiasmata. A base number 
of r — 6 is not known for any of the tribes within the Caesal- 
pinioideae (Darlington and Wylie, 1956). However, since a 
base number of x = 12 is common for many genera of the Caesal- 
pinioideae it might be conjectured that the number x = 6 for 
Krameria is but a lower base for the subfamily as a whole. Con- 
sideration of chromosome morphology proves more instructive. 
Meiotic chromosomes from a wide selection of Cassia species 
(Turner, 1956; H. 8. Irwin, unpublished) are consistently small, 
usually showing only two terminalized chiasmata at metaphase. 
This is also true of the known meiotic chromosomes of species 
examined in the tribes Bauhineae and Eucaesalpinieae. By com- 
parison, meiotic chromosomes of Krameria are 10-40 times as 
massive as those of the Cassieae thus far examined. 

Unfortunately, Krameria can not be compared with the Poly- 
galaceae since chromosomal information on the family is scanty. 
The only established base number for the family is x = 7 and 
this from a single mitotic count on Bredemeyera colletioides (Covas 
and Schnack, 1946). 

Though the cytological evidence available at present does not 
permit one to judge phyletic alternatives, at least chromosomal 
comparisons of Krameria with members of the subfamily Caesal- 
pinioideae seem to negate any close relationship with taxa of this 
group. In view of this negation, particularly as concerns its 
affinity with the Cassieae, the genus Krameria, even if related to 
the Caesalpinioideae on phyletie grounds, seems deserving of 
supra-tribal rank, if cytological evidence is considered along with 
that of wood anatomy, floral morphology, etc. 

“Problem” taxa such as Krameria make present-day taxonomy 
the exciting field it is. One never knows how new evidence will 
affect the taxonomic scales. Indeed, it stimulates the taxonomist 
to look to other fields for additional weights that might affect 
the balance. Thus floral morphology, anatomy, palynology, 

5 These authors (p. 148) list 6 as one of several base numbers for Cassia, but this is not borne 
out by a reference to their listed counts. Senn (1938) gives a count of n = 6 for Cercis canaden- 
sis, but Baldwin (1939) reported counts of 2n = 14 for several collections of the species. In 


view of the drawing accompanying Senn's report (p. 183), which appears to show two clumped 
chromosomes drawn as one, the report of n = 6 for this species should be considered erroneous. 


1958] Turner,—Chromosome Numbers in Genus Krameria 105 


3 


Figures 1-4. Camera lucida drawings of the meiotic chromosomes in Krameria spp.—Fig. 1 s 
K. lanceolata, chromosomes closely packed at metaphase I.—Fig. 2. K. lanceolata, metaphase 
of division II.—Fig. 3. XK.ramosissima, metaphase of division II.—Fig. 4. K. grayi, anaphase 
of division II (only one half of quartet shown). (X ca 1400). 


biochemistry, cytology, ete., must necessarily bring us closer to 
the truth, since the total attributes of a group of organisms will 
more nearly reflect their relationships than will the characters 
from any one field when considered alone. 

If all taxonomie problems were obvious and merely resolved 
themselves to eataloguing always discreet, easily placed entities, 
then many of us would long since have lost interest in the tabula- 
tion and turned to other fields. 


106 Rhodora [Vor. 60 


LITERATURE CITED 


ABRAMS, L. 1944. Krameriaceae. Illustrated flora of the Pacific states 2: 
627—628. 

BarnpwiN, J. T., JR. 1939. Chromosomes from leaves. Science 90: 240. 

—————-—-, AND B. M. SrkEsE. 1957. Phoradendron flavescens: chromo- 
somes, seedlings, and hosts. Amer. Jour. Bot. 44: 136-140. 

Benson, L. 1957. Plant Classification. 688 pp. D. C. Heath and Company, 
Boston. 

Benson, L., anb R. A. Darrow. 1954. The trees and shrubs of the South- 
western deserts. 437 pp. University of Arizona Press, Tucson. 

BENTHAM, G., AND J. D. Hooker. 1862. Polygalaceae. Genera Plantarum 
1: 134-140. 

Brirron, N. L. 1930. Krameriaceae. North Amer. Flora 23: 195-200. 

CAPITAINE, L. 1912. Étude analytique and phytogéographique du groupe 
des Légumineuses. 496 pp. Paris. 

CuopaT, R. 1890. (without title). Archives des Sciences Physiques et 
Naturelles Ser. 3. 24: 405—499. 

Covas, G., AND B. ScuNACK. 1946. Numero de cromosomas en antofitas de 
la region du Cuyo (Republica Argentina). Rev. Arg. Agron. 13: 153-1606. 

CnoNcuisT, A. 1957. Outline of a new system of families and orders of 
dicotyledons. Bull. Jard. Bot. de l'État Bruxelles 27: 13-40. 

DARLINGTON, C. D., and A. P. WvurE. 1956. Chromosome atlas of flowering 
plants. 519 pp. Macmillan Company, New York. 

EnpTMAN, G. 1944. The systematic position of the genus Diclidanthera. 
Bot. Notiser 1944: 80-84. 

HALLER, H. 1912. L'origine et le système phylétique des angiospermes. 
Arch. Néerl. Sci. Exact. Nat. Ser. III B, 1: 146-234. 

HkrMscH, C. 1942. Comparative anatomy of the secondary xylem in the 
“Gruniales” and *"Terebinthales". Lilloa 8: 83-198. 

HvrcnuiNsoN, J. 1926. The families of flowering plants. I. Dicotyledons. 
328 pp. London. 

Kunz, M. 1913. Die systematische Stellung der Gattung Krameria unter 
besonderer Berücksichtigung der Anatomie. Beihefte Botanisches Cen- 
tralblatt 30: 412-417. 

Senn, H. A. 1938. Chromosome number relationships in the Leguminosae. 
Bibliographia Genetica 12: 175-337. 

SMALL, J. K. 1903. Flora of the Southeastern United States. New York. 

TAUBERT, P. 1894. Leguminosae. In Engler and Prantl, Die Natürlichen 
Pflanzenfamilien IHI’: 70-396. 

Turner, B. L. 1956. Chromosome numbers in the Leguminosae. I. Amer. 
Jour. Bot. 43: 577—581. 


Waterfall,—Genus Physalis in N. America 107 


A TAXONOMIC STUDY OF THE GENUS PHYSALIS 
IN NORTH AMERICA NORTH OF MEXICO! 


U. T. WATERFALL 


LiNNAEUS founded the genus Physalis in 1753 with nine species, 
two of which are now usually referred to Withania. Five of the 
remaining seven are found within the area of the present study, 
although one, P. Alkekengi, is known only as a horticultural 
species, or as an escape from cultivation. In the second edition 
of Species Plantarum (1762) Linnaeus added two more species, 
P. pensylvanica and P. peruviana. The former does not occur in 
Pennsylvania and perhaps is not found in North America. In 
any event, it is considered to be a synonym of P. viscosa. P. 
peruviana is sometimes cultivated. Perhaps it escapes, but, if so, 
it is rarely collected. Pre-Linnean authors were familiar with 
the genus under the names Physalis, Solanum and Alkekengi as 
indieated in the Linnean references. Philip Miller (1768) de- 
scribed two species coming within the scope of this study. One 
of them, P. virginiana, is here interpreted as it has been for the 
last sixty years, although the application of the name is by no 
means certain. 

Michaux (1803) described P. lanceolata and P. obscura, the 
latter consisting of var. glabra and var. viscidopubescens. Nees 
(1831) described P. heterophylla, P. lanceifolia and P. Linkiana. 

P. crassifolia, the common species of southwestern desert re- 
gions, was described by Bentham (1844). "Thomas Nuttall, in 
publications hereinafter enumerated under the species concerned, 

! This paper is based on a dissertation submitted in partial fulfillment of the requirements fo" 
the degree of Doctor of Philosophy at the University of Oklahoma under the guidance of Dr- 
George Goodman, Professor of Plant Sciences. The author is grateful to him, and to the other 
members of his committee, for their valuable suggestions. 

He is also thankful to the curators of several herbaria whose loans of all, or selected parts, of 
their valuable collections of Physalis made this study possible. They have loaned 8090 sheets 
of Physalis and related genera. Of this total, there are 5716 sheets of Physalis included in the 
present study. Many of the remainder were collections from other areas which materially 
aided in the formation of species concepts. These herbaria, listed by the standardized abbre- 
viations published by Lanjouw and Stafleau (1954), are: ARIZ, COLO, DUKE, GH, KANU, LIL, 
MICH, MO, NY, OKL, OKLA, P, PH, RM, SMU, TEX, UARK, UC and wis. 

Thanks are also due the librarians of both the University of Oklahoma and Oklahoma State 
University (formerly Okla. A. & M. College) for securing photostats of publications unavailable 
locally, and the Arnold Arboretum of Harvard University for the loan of photographs of Lin- 
nean types, as well as to Dr. Albert Delisle formerly of the University of Notre Dame for photo- 


graphing certain types of species described by Greene. 
This study was made possible, in part, by a grant from the Southern Fellowships Fund. 


108 Rhodora [Vor. 60 


described P. angustifolia, P. longifolia, P. mollis, P. pumila and 
P. Walteri. Dunal (1852) described P. nyctaginea, now generally 
referred to P. heterophylla. 

In his study of the North American species, Asa Gray (1875) 
described P. Fendleri, P. hederaefolia and P. Wrightii. Rydberg 
(1896), in the last general study of the genus, described P. ciliosa, 
P. comata, P. macrophysa, P. neomexicana, P. rotundata and 
P. versicolor. 

Following this, as a result of increased botanical activity, Mohr 
(1899) described P. monticola; P. rigida was described by Pollard 
and Ball (1900), and P. polyphylla by Greene (1900). P. 
missouriensis and P. subglabrata were proposed by Mackenzie and 
Bush (1902). Rydberg added P. floridana, P. pendula and P. 
sinuata in Small’s Manual (1903). Aven Nelson (1909) proposed 
Quincula lepidota and P. genucaulis. P. caudella, described by 
Standley (1937) from Chihuahua has been found in southern 
Arizona, and these collections usually have been identified as 
P. lanceolata. 

Margaret Y. Menzel (1951) has published an exploratory sur- 
vey of the eytology and geneties of many of the species of our 
area, as they were interpreted by Rydberg, and various collectors 
who followed that author's treatment. Her work is valuable 
because it shows that the observed variation may indeed be 
correlated with cytologic and genetic differences. 


TAXONOMIC CHARACTERISTICS 


The stability of taxonomic characteristics apparently has been 
much disturbed by the long-continued distribution of various 
species by man as esculents. Some of the species, such as P. 
ixocarpa and P. peruviana, are still so used. No doubt many of 
the species have been used by aborigines of many parts of the 
world at one time or another. This introduces the possibility of 
gene interchange between taxa that would otherwise not be con- 
tiguous. Furthermore, man’s activities in producing disclimaxes 
provide habitats in which individual plants, or populations, of 
narrow ecological amplitude may survive. Such ecological niches 
might not have existed otherwise. It may be significant that 
many collections of Physalis are made in such disturbed habitats. 
Somewhat similar situations are admirably discussed by Edgar 
Anderson (1949). 


1958] Waterfall, —Genus Physalis in N. America 109 


The following discussion concerns the principal morphological 
characteristics which have been used in the taxonomy of the 
genus. 

roots. Both annual and perennial species occur in the genus. 
The former have fibrous root systems, or sometimes taproots. 
The latter are usually woody-based or rhizomatous. The annual 
vs. perennial characteristic has been used much in the past, 
beginning with Linneaus, to divide the genus into two main 
divisions. Since most herbarium specimens do not show the 
underground parts, this system is not a particularly convenient 
one. More dependence has been placed on other struetures in 
the following treatment. 

STEMS. All of our species have herbaceous stems, although in 
Mexico and Central America shrubby ones may occur. They 
may perennate from a woody caudex in such species as P. crassi- 
folia. of the desert areas of southwestern United States. Several 
of them grow from rhizomes, which may be cord-like, near the 
surface, and are often present in herbarium collections, as P. 
arenicola of sandy areas of Georgia and Florida, or thick, deeply 
buried and seldom collected, as the wide-ranging P. heterophylla 
and P. virginiana. The stems may be erect to prostrate and 
simple to much branched. An extreme in branching is found in 
P. crassifolia which, as reported by Jepson (1925), forms plants 
“1-3 ft. broad and 15-1 ft. high." 

LEAVES. The leaf blades are usually ovate to ovate-lanceolate, 
but they may be reniform, as are rarely some of the lower leaves of 
P. hederaefolia, or narrowly linear as in P. angustifolia. Petioles 
may be longer than the blade, or the blades may taper into short, 
winged petioles. The leaves are normally alternate, but some- 
times appear to be opposite as in P. viscosa var. Elliottii which 
often has two leaves at a node, particularly in the upper part of 
the stem. 

Leaf shape has often been used as a basis for establishing, or 
characterizing, taxa, as is indicated by such names as latifolia, 
hederaefolia, angustifolia, heterophylla, lanceifolia, integrifolia, 
crassifolia, spathulaefolia and longifolia. No doubt in some in- 
stances, as in P. angustifolia with its long linear leaves, the em- 
phasis on leaf-form is justified. However, the size, shape and 
margins of leaves are extremely variable characteristics in groups 
that appear to be natural populations. One approach to the 


110 Rhodora [| Vor. 60 


understanding of this variability is to study a species that is well- 
delimited otherwise. An excellent example is P. lobata, a species 
so distinct that it has been considered a separate genus by such 
taxonomists as Rafinesque and Rydberg. Here the leaf shape 
may vary from narrowly ovate to linear-oblong, and the margins 
from pinnatifid to entire. Although the variation in leaf mar- 
gins appears to be at an extreme here, the variability of leaf shape 
can be duplicated, and the variability in margins approached, in 
several other natural populations. 

Under such conditions it would seem hazardous to establish 
species or varieties based on these characteristics; such a pro- 
cedure should be followed only when these features are correlated 
with other morphological characteristics, or with geographic dis- 
tribution. The author has utilized these criteria in maintaining 
P. viscosa var. spathulaefolia of the Texas gulf, and P. viscosa var. 
maritima of the southeastern seacoast. Here the extremes are 
distinet, but variation makes some collections difficult to place. 
Measurements of many specimens show the leaves of the former 
to be narrower than the latter, as indicated later in the keys to the 
species. However, if these features are used with the idea of 
either matching specimens, or describing new species, only con- 
fusion can result. 

vESTITURE. The indument varies from hairs that are stellate 
or variously branched, to long jointed hairs and short hairs, 
capitate or sessile glands, or small crystalline vesicles as in P. 
lobata. Even the seemingly glabrous species usually have a few 
trichomes of some kind, at least on the younger parts. Fre- 
quently two or more kinds of trichomes are intermixed. 

Such names as viscosa, mollis, comata, pubescens, hirsuta, 
cinerascens, ciliosa, villosa, pruinosa and subglabrata indicate the 
consideration that authors have given to indument in the past. 
The procedure seems to be partly justified. Surely the “stellate” 
populations are related. But if one attempts to distinguish taxa 
on the basis of the density or the size of the stellate hairs, caution 
should be exercised. In this study P. viscosa var. mollis has been 
segregated from var. cznerascens partly by this characteristic, but 
the latter taxon is extremely variable within itself in this respect. 
In P. heterophylla many of the variations in vestiture seem to be 
so little correlated, either with other characteristics, or with 


1958] Waterfall,——Genus Physalis in N. America 111 


geographie distribution, as to be unusable to distinguish even 
varieties. On the other hand, in P. virginiana, sens. lat., the 
correlations are such that they are of value in helping to establish 
geographie varieties. In the P. angulata-P. pubescens series in- 
dument is also of taxonomic significance, the villous P. pubescens 
usually having abundant multi-cellular hairs, P. angulata having 
a few short ones. 

COROLLAS. The shape, color and spotting of the corollas have 
been considered of taxonomic significance. The shape varies 
from funnel-form-campanulate to rotate with the limb reflexed. 
The corolla is plicate, and is truncate with the exception of 
P. Alkekengi in which the lobes are separated by short sinuses. 
The shape of the corollas may be of taxonomic significance. 
Since the characteristic shape is attained for only a short time in 
the full sun, the application of this criterion is of limited value. 
Several species such as P. lobata, P. Wrightii, P. hederaefolia and 
P. crassifolia have corollas that are either rotate, or have a re- 
flexed limb when fully open. Since the corollas are seldom fully 
open, the author has usually used a linear measurement for com- 
parative purposes where such usage seemed desirable. 

Color of the corolla has been used to help characterize P. lobata, 
which is our only species with a bluish, or violet, corolla. Other- 
wise the presence, and sometimes the color, of five spots on the 
limb of the yellowish corolla near its base has been found useful. 
The majority of the species either have distinct, dark spots 
present, or they have none that are noticeable in herbarium 
specimens. A few taxa such as P. hederaefolia var. Fendleri and 
some of the maritime varieties of P. viscosa have spots which are 
only a little darker than the rest of the corolla. 

STAMENS. Size and color of the anthers are of taxonomic value. 
With a few exceptions, large anther size (measurements given in 
the keys) and thick filaments are correlated with our perennials. 
Small anther size and slender filaments are usually found in our 
annual species. In some taxa anther color is of significance. So 
many of the annual species have blue anthers that the yellow 
anthers of P. missouriensis attract attention. In others, such as 
P. heterophylla and P. virginiana var. virginiana, yellow or bluish 
tinged anthers seem to occur without much significance. How- 
ever, P. virginiana var. subglabrata and var. sonorae may be dis- 


112 Rhodora [ Vor. 60 


tinguished by the bluish anthers of the former and the yellow 
anthers of the latter even when their other characteristics overlap. 

A peculiarity of P. crassifolia and its var. versicolor is the 
presence of a few long jointed hairs on the filaments. 

FLOWERING CALYX. The relative depth to which the calyx 
lobes are divided may be of value, as it is in helping to separate 
P. angulata var. angulata from var. pendula. 

PEDUNCLE. The length of the peduncle, both flowering and 
fruiting, may be of taxonomic significance. In fruiting material 
of P. ixocarpa and P. virginiana var. subglabrata, some specimens 
of which may resemble each other, the very short fruiting 
peduncle of the former will serve to separate the two. Among the 
southwestern desert species, P. hederaefolia and its relatives may 
be separated from P. crassifolia and its relatives by the short 
flowering peduncle of the former. Although of lesser significance 
in the P. angulata complex, it may be used, in conjunction with 
the size of the fruiting calyx, to help separate P. angulata var. 
angulata from var. pendula and var. lanceifolia. 

FRUITING CALYX. The calyx greatly enlarges with the matur- 
ing fruit, usually being much inflated around it. In some popula- 
tions the size and shape seem to be constant, and characteristic 
enough to be taxonomically usable. In P. pubescens and its 
relatives there is present a distinctly five-angled fruiting calyx. 
A population in southern Arizona is proposed as a new species, 
easily recognized by its unusually broad, sharply-angled fruiting 
calyx. In the P. angulata series, P. angulata var. angulata has a 
larger fruiting calyx than either var. pendula or var. lanceifolia. 

In other populations the size and shape of the fruiting calyx 
seem to be either quite variable, or the extremes occur 
sporadically. The present author believes that the large-calyx 
form described as P. macrophysa is a more or less sporadically 
occurring form of P. virginiana var. subglabrata, although it also 
may be found in intergrades with var. sonorae (P. longifolia). 
Specimens with large fruiting calyces also appear in P. virginiana 
var. virginiana and in some phases of P. viscosa. 

'The length of the lobes of the fruiting calyx was considered 
characteristic enough by Standley (Lle.) to call a new species 
P. caudella. 

The writer has not found the indentation at the base of the 


1958] Waterfall, —Genus Physalis in N. America 113 


fruiting calyx to be of much taxonomie value. Considerable 
variation may occur on the same plant. Of course if the calyx is 
nearly filled by the berry, it will be little invaginated. 

OTHER CHARACTERISTICS. The style has not been used to any 
extent. In P. lobata it is distinctively curved near the base and 
bent to one side. The more or less reniform, punctate to reticu- 
late seeds are very similar in most of the species. Differences 
seen in preliminary study appear to be bridged by many inter- 
mediates when a large series is examined. The backs of the seed 
of P. lobata are rather crenate or rugose. 


GENERIC RELATIONSHIPS 


The genus Physalis is studied here in its more or less conven- 
tional conception, including those members of the Solanaceae 
having a funnelform or campanulate to rotate, or rotate-reflexed 
corolla, longitudinally dehiscing anthers, and a berry, or berry- 
like fruit, enclosed in an enlarged and usually inflated calyx. It 
excludes both Margaranthus, with its urceolate corollas and very 
Physalis-like fruiting calyces, and also Chamaesaracha with a 
corolla very similar to some species of Physalis, but with a fruit 
very closely invested by the enlarging calyx. 

When not in flower, Margaranthus could hardly be dis- 
tinguished from moderately small-fruited species of Physalis. 
The rather tightly investing fruiting calyx of Chamaesaracha can 
be matched, or is approached, in some specimens of certain species 
of Physalis such as P. ixocarpa. A peculiar situation is found in 
Chamaesaracha where C. grandiflora, originally described as Phy- 
salis by Hooker, and a related species, C. nana, have seeds very 
similar to the punctate or minutely reticulate seeds of Physalis, 
while the other species of the genus have rather strongly alveolate 
surfaces. 

Possibly both genera should be included in Physalis. This 
would make PAysalis an inclusive genus, similar in its concept to 
Oenothera as delimited by Munz and other conservative taxono- 
mists. In that genus a number of subgenera, regarded as genera 
by some authors, are bound together by flower similarities despite 
their differently shaped fruits. In Physalis, under this broad 
concept, the distinctive fruiting calyx would hold together sub- 
genera differing in corolla structure. In Chamaesaracha the dis- 


114 Rhodora [Vor. 60 


tinctiveness of the fruiting calyx becomes progressively less 
evident. The difficulty here would be in finding a stopping place 
short of including the whole genus. The author prefers making a 
more intensive study of the problem before proceeding with such 
action. 

Since there is the possibility of creating distinctive subgenera 
as outlined in the preceding paragraph, it seems preferable not to 
formally place the species here treated into subgenera or sections, 
but to defer this action until not only species of Physalis from 
other areas, but also related taxa can be studied. 


TAXONOMY 

PHYSALIS L., Species Plantarum 1: 182. 1735; Alkekengi ‘Tourn. ex 
Hall, Enum. Stirp. Helv. 2: 508. 1742; Herschellia Bowdich, Excurs. 
Mader. 159. 1825. Quincula Raf. Atl. Journ. 145. 1832; Alicabon Raf., 
Sylva Tellur. 56. 1838; Pentaphiltrum Reichb., Das Herbarienbuch 121. 
1841; Boberella Krause, in Sturm, Fl. Deutschl. ed. 2 (10): 54. 1903. 

Plants annual or perennial with herbaceous stems, some having woody 
caudices, others with short to elongated rhizomes; leaves usually broadly 
ovate to linear, alternate or sometimes two at a node; vestiture various in 
kind and quantity, including short hairs, long jointed hairs, stipitate or 
sessile glands, or with hairs variously branched to stellate; corollas plicate, 
campanulate to rotate with the limb reflexed; corolla color usually some 
shade of yellow with, or without, five darker spots near the base of its limb, 
sometimes blue; flowers usually solitary in the axils of the leaves, some- 
times on foreshortened axillary branches causing them to appear to be in 
axillary fascicles; calyx united, its lobes distinct for a little over one-half to 
about one-fourth of its length; calyx lobes ovate-deltoid to narrowly lan- 
ceolate, sometimes acuminate; calyx enlarging with, and usually inflated 
around, the maturing fruit; fruit a two-carpellate many- to few-seeded 
berry, sometimes rather dry; style more or less filiform, usually expanding 
somewhat at its summit into a slightly capitate, but sometimes nearly 
truncate, stigma; stamens five, their filaments attached near the base of 
the corolla tube; anthers ovate-oblong to linear-oblong, dehiscing by lateral 
slits, yellow or bluish in color; filaments varying from nearly as wide as the 
anthers, and sometimes clavate, to slender and filiform. 


(To be continued) 


1958]. Fell, —New Illinois Carex Records 115 


NEw Ivurnois Carex Recorps.'—Collections of the following 
species of Carex have been deposited, as indicated, in Illinois State 
Museum (sm), University of Illinois (1v), University of Wisconsin 
(wu), and Rockford College (rc), herbaria. 

Carex stenophylla Wahl. var. enervis (C. H. May) Kiikenth. 
(C. eleocharis Bailey) ranges normally west and northwest from 
Iowa. It has not been reported east of the Mississippi River. 
In the spring of 1957 it was found on a dry gravel bluff prairie in 
Greater Rockford Airport south of Rockford, Winnebago County, 
in an area that was a part of Camp Grant during World Wars I 
and II. Here it grows in nearly pure stands in several patches 10 
to 20 feet in diameter, blooming at the same time as C. pensyl- 
vanica Lam. var. digyna Boeckl. with which it is associated. It 
ceases growth by the first of July at which time the slender rhi- 
zomes and stolons have reached a length of 21% to 3 inches. Thus 
it seems likely that seeding took place during World War I. Col- 
lection numbers are: 57-9 (sM, IU, WU, RC); 57-68 (SM, IU, RC); 
57-157 (SM, IU, WU, RC); 57-248 (SM, IU). 

Carex praegracilis W. Boott, another western species, is credited 
in the manuals to northern Michigan. There are no other records 
of its occurrence east of the Mississippi River. In 1951, it was 
found at Greater Rockford Airport and since then in a number of 
places on the gravel bluff prairie in the area, on a roadside near 
Perryville six miles from the Airport and in DeKalb County near 
Kirkland on a railroad right-of-way, 15 miles distant. Neither of 
these places are on a direct line of travel from the Airport. All 
are in prairie situations. The strong rhizomes grow rapidly but 
definite patches are not formed so the probable length of time that 
it has been established cannot be determined. The World War I 
idea of introduction would not apply to the DeKalb County sta- 
tion. Station locations and collection numbers are: Winnebago 
County; (Rockford Airport) 51—114 (sw); 52-356 (sm); 54-420 
(wu); 57-158 (1v); (Perryville) 55-440 (wv, rc); 56-73 (1v). 
DeKalb County (Kirkland) 53-371 (sm); 55-141 (wv); 56-91 (1v). 

In this location these plants bear seed sparingly but C. prae- 
gracilis is well dispersed and seems to be well established. C. 
stenophylla has persisted for 40 years and has spread. It is in a 
favorable habitat which is of a type common in Winnebago 
County. The airport authorities have agreed that this gravel hill 


1 A contribution from the Evelyn I. Fernald Memorial Herbarium of Rockford College. 


116 Rhodora [Vor. 60 


prairie will not be disturbed unless it is needed for an essential 
operation of the airport, which is not likely. For these reasons it 
seems probable that these carices will become a part of our flora 
with Paspalum  stramineum, | Ratibida | columnifera, Froelichia 
gracilis, Artemisia dracunculoides, and others from the west which 
are found in the same area.—EGBERT W. FELL, 


NELUMBO LUTEA IN Essex County, MassAcHUSETTS.— lhe 
only known station for Nelumbo lutea (Willd.) Pers. in Essex 
County was the Devil's Dishfull in West Peabody where it was 
apparently introduced at some time between 1880 and 1918. It 
soon became so abundant that the pond became known as Lotus 
Pond and some well-intentioned but misinformed person erected 
a sign stating that the pond was the only place in the United States 
where the Sacred Lotus occurred. Nelumbo was still common there 
in the fall of 1953, when I collected a specimen. In August, 1957, 
I put my boat in the pond and covered the area thoroughly collect- 
ing aquatic plants. While Nuphar variegatum and Nymphaea 
odorata were as common as in the past there was no trace of 
Nelumbo. I ean think of no valid cause to explain its disap- 
pearance.— STUART K. HARRIS, DEPT. OF BIOLOGY, BOSTON UNI- 
VERSITY. 


CABOMBA CAROLINIANA IN Essex County, MASSACHUSETTS. — 
When collecting aquatics in Fosters Pond, Andover, Massachusetts 
in June, 1957, I found that the most common plant there was 
Cabomba caroliniana Gray. This is the first station known in 
Essex County. Since Professor A. S. Pease, who has an uncanny 
ability to spot interesting plants, collected on the pond in 1903 
and did not find Cabomba, it seems safe to assume that it must 
have been introduced there since that date. Probably Cabomba 
is more widely introduced in Massachusetts than collections indi- 
cate. There are only two sheets in the herbarium of the New 
England Botanical Club from this state other than my collection, 
24 June 1957, 12997. However, Cabomba is abundant in Muddy 
River in Boston’s Fenway but no specimen appears in the Club 
herbarium.—Struarr K. HARRIS, DEPT. OF BIOLOGY, BOSTON 
UNIVERSITY. 


Volume 60, no. 711, including pages 61-88, was issued 25 April, 1958, 


Douova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS l 
RALPH CARLETON BEAN 
CARROLL EMOR Y WOOD, JR ( 
IVAN MACKENZIE LAMB 


Associate Editors 


Vol. 60 May, 1958 No. 713 
CONTENTS: 
Generic Considerations Concerning Carphephorus, Trilisa and 
Litrisa (Compositae). Charles W: James.................. 117 
Chromosome Races in the Chrysanthemum Leucanthemum Com- 
D ONSE PREIS LL eiu re CET 122 
Notes on the Distribution of Ohio Compositae: I. Heliantheae, An- 
themideae. Hobert W. Long.: otasidan cee cece eee nnn 125 
A Taxonomic Study of the Genus Physalis in North America North 
of Mexico. U. T. Waterfall (continued from p. 114)......... 128 
Further Notes on the Illinois Flora. Robert A. Hvers............ 142 


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Vol. 60 May, 1958 No. 713 


GENERIC CONSIDERATIONS CONCERNING 
CARPHEPHORUS, TRILISA AND 
LITRISA (COMPOSITAE) 


CHARLES W. JAMES 


Carphephorus Cass. (4 species), and Trilisa (Cass.) Cass. (2 
species) and the monotypic Litrisa Small (merged with Trilisa by 
Robinson, 1934), are closely related genera in the Eupatorieae. 
They are allied to Liatris Schreb. and Garberia Gray with which 
they form a seemingly natural segregate of the subtribe 
Kuhniinae. Primarily of the Coastal Plain, they are known only 
from the southeastern United States; southeastern Virginia to 
south Florida and westward into eastern Louisiana. All of the 
seven species appear distinctive and present relatively little 
variation. The present problem is one of generic limits, the basis 
of which is presented in the following historical account. 

In describing Carphephorus, Cassini (1816) stated that it dif- 
fered from Liatris in that the receptacle was provided with pales 
and the pappus bristles were non-plumose. Later (1818), he 
recognized Trilisa as a subgenus of Liatris, as typified by Liatris 
odoratissima. The rank of subgenus was considered sufficient 
for this taxon because there was observed in Liatris a short- 
plumose (barbellée) pappus which was intermediate between true 
Liatris whose pappus was long-plumose (barbée), and Trilisa 
whose pappus was barbed (barbellulée). He further stated that 
Trilisa had the greatest affinity with Carphephorus, from which 
it differed by the absence of pales. Although he directly pro- 
ceeded to point out that a few pales were occasionally observed 
even in the subgenus Trilisa. In 1820, Cassini raised Trilisa to 
generic status without further discussion. Not until 1828, how- 


118 Rhodora [ Vou. 60 


ever, were any species actually transferred to it. At this time 
Cassini cited Liatris odoratissima and Liatris paniculata of Will- 
denow (1803)! as species of Trilisa and summarized his previous 
comments on these genera. 

It is Cassini’s remark (apparently overlooked, subsequently) 
of the occasional presence of some pales in T'rilisa odoratissima 
that has prompted the question of the generic limits in Carphe- 
phorus and Trilisa, and also Litrisa. The last named “genus” 
has only one species, Litrisa carnosa Small, an endemic of east, 
central Florida. In describing it, Small (1924) wrote, “Tech- 
nically it is most closely related on the one hand to T'rilisa, by its 
involuere, and on the other, to Carphephorus, by its chaffy re- 
ceptacle." It is implied here, as has been customary, that Trilisa 
has a naked receptacle. After the publication of Small’s manual 
(1933), Robinson (1934) noticed that Small contradicted himself 
by keying out Litrisa as having “Receptacle naked." In a study 
of the type collection, as well as other material, Robinson made 
several sketches, one of which bears the annotation, “No scales 
on disk." Since he then proceeded to transfer Litrisa carnosa to 
the genus Trilisa, rather than Carphephorus, the implication is 
again that Trilisa lacks pales. Presumably, if Robinson had 
seen any pales in Litrisa carnosa, he would have transferred it to 
Carphephorus, for he was not impressed by the differences in the 
involucres. 

Upon examination of all of the species of the genera under con- 
sideration, I conclude that pales may be borne in any one of the 
species. They are most abundant in C. pseudo-liatris and C. 
corymbosus. Their number also varies, as might be expected, 
with the size of the heads. Since the heads are typically smaller 
in the two Trilisa species and in Litrisa (involucres ca. 4-5 mm. 
high), than in Carphephorus (involucres ca. 6-10 mm. high), the 
number of pales per head as a primary generic character would 

1 Willdenow took both epithets, ‘‘odoratissima’’ and “‘paniculata’’ from Walter (1788), who 
had employed them with *"Anonymos." However, Michaux, also in 1803, transferred these 
"Anonymos' species of Walter to Liatris. According to Schubert (1942), Michaux's Flora 
preceded this particular volume of Willdenow’s. Consequently, the citation of these binomials 
should be: Trilisa odoratissima (Michx.) Cass. and Trilisa paniculata (Michx.) Cass. 

2 The presence of pales in these genera is unique in the tribe Eupatorieae. Bentham (1873) 
says, ‘‘Receptaculum plus minus paleaceum in Decachaetae, Alomiae, et Agerati speciebus 
paucis etin Carphephorus. They are also present in Hartwrightia Gray ex S. Wats. Further- 


more, they are little, if at all, specialized; the peripheral ones, particularly, are very similar to 
the phyllaries, with which they have been considered homologous. 


1958] James,—Generie Considerations 119 


seem questionable. The pales are also deciduous, a factor per- 
haps accounting for discrepancies in determining their presence? 
or absence. 

The only other known morphological basis for these genera is 
found in the involucre. In Carphephorus, the phyllaries are well- 
imbricated in 3-6 series; the involucre 6-10 mm. high. In 
Trilisa, the phyllaries are in 1-2(-3) series, scarcely, if at all, 
imbricate, the involucre 4-5 mm. high. However, in Litrisa, the 
phyllaries are well-imbricated, but only in 2-3 series and the 
involuere is 4-5 mm. high. 

Cytological studies (Gaiser, 1954) in these genera have yielded 
no evidence for the maintenance of Trilisa or Litrisa as distinct 
from Carphephorus. In all of the species 2n = 20. ‘One karyo- 
type is believed to be common to these two species [T'rilisa 
paniculata and Trilisa odoratissima] and it has been found to be 
indistinguishable from that of Carphephorus.’’ Concerning 
Litrisa carnosa, “The number (2n = 20) and approximately the 
same kinds of chromosomes as found in the other two species 
[of Trilisa] were sketched from cells not adequate for photog- 
raphy. However, lacking sufficient material for careful studies, 
the karyotype of this species cannot be included at this time.” 
The only other genera in the subtribe Kuhniinae in which n = 10? 
are Garberia and Liatris, both of which have long been suspected 
of being closely related to the genera in question by their mor- 
phological similarities. Neither of these genera has been reported 
to have pales. The monotypic Garberia, represented by G. 
heterophylla (Bartr.) Merrill & F. Harper (G. fruticosa (Nutt.) 
Gray), is endemic to the sand scrubs of central Florida and is the 
only woody member having n = 10. Also, its karyotype is dis- 
tinctive. Although karyotypes have not been determined for 
all of the species of Liatris, some intra-generic variation in karyo- 
type has been detected. Certainly there is as much variation, 
morphological as well as cytological, in the one genus Liatris (cf. 
Gaiser, 1946; 1949; 1950) as there is in the whole Carphephorus- 
Trilisa-Litrisa complex. 

As it now stands, the primary basis for Trilisa (including 
Litrisa, as Robinson did) seems to rest entirely on the size of the 


3 Other basic numbers in the subtribe are 9 and 11; only the South American genus, Kanimia, 
has not been examined cytologically. 


120 Rhodora [Vor. 60 


involucres. If, however, Litrisa carnosa were transferred to 
Carphephorus, the primary generic distinction could then be based 
on whether or not the phyllaries were imbrieate. The latter 
character would appear to segregate the species much more 
naturally. This apparently was R. M. Harper's view, also, for 
he collected T'rilisa carnosa before it was described (St. Lucie Co.: 
flat pine woods about 2 mi. w. of Fort Pierce, Fla., 23 Aug. 1923, 
GH), and annotated it **Carphephorus (?) n. sp." This same char- 
acter has been used as a primary basis for distinguishing 
Brickellia from Kuhnia (cf. Robinson 1913; 1917), also in the 
Kuhniinae, but having n = 9. Shinners (1946) considered this a 
very weak basis, but in this case was able to point out additional 
reasons to justify the continued recognition of those genera. 
In the present and somewhat comparable case, supplementary 
characters, if any, have not been found. But regardless of 
whether or not Trilisa is to be maintained as a genus, it appears 
best segregated at the present time on its non-imbricate phyl- 
laries. Since the phyllaries of Litrisa are imbricate as in Car- 
phephorus, it is proposed that Litrisa carnosa Small be transferred 
from Trilisa to Carphephorus. This action would be in agreement 
with MeVaugh's (1945) recommendation six on the generic dis- 
position of species having affinities with two or more genera. 
“Any segregate genus should be sharply delimited; that is, any 
species which is intermediate in one or more respects toward a 
more inclusive genus should be relegated to the latter. The re- 
tention of the anomalous species in the more inclusive genus will 
change its limits, if at all, but very slightly, and only in this way 
can the segregate genus be precisely defined." In this case, the 
“segregate genus" would be Trilisa, the ‘more inclusive genus"— 
Carphephorus, and the “anomalous species’’—Litrisa carnosa. 


Carphephorus carnosus (Small) James, comb. nov. based on Litrisa 
carnosa Small, Bull. Torr. Bot. Club 51: 392. 1924. Type: Small 10658, 
Istokpoga Prairie, east of Sebring, Florida, 31 Aug. 1922 (wy); photograph 
of type (au). Trilisa carnosa (Small) Robinson, Contrib. Gray Herb. 
104: 49. 1934. 

This species is endemic to the seasonally wet, low, sandy pine lands in 
east central to southern Florida. It is known from Brevard (anu), Char- 
lotte (FLAs, GH), DeSoto (rtas), Highlands (rtas, an), Martin (FLAs, GH), 
Okeechobee (FLAS, GH), Orange (FLAS, GH), Osceola (FLAS, GH), Polk (FLAS) 
and St. Lucie (aH) Counties. 


1958] James,—Generie Considerations 121 


KEY TO THE SPECIES 


a. Phyllaries imbricate (in (2-)3-6 series), densely pubescent 
with eglandular trichomes or glabrous and erose-ciliate; 
involucre 4-10 mm. high......................00. Carphephorus Cass. 
b. Phyllaries (at least the inner) glabrous, the margins erose- 
ciliate, eglandular, obtuse. 
c. Stems pubescent.................. 1. C. corymbosus (Nutt.) T. & G. 
c. Stems glabrous.................. 2. C. bellidifolius (Michx.) T. & G. 
b. Phyllaries with eglandular trichomes and colorless resin 
atoms (sometimes few), acute or apiculate. 
d. Basal leaves long, needle-like............... 3. C. pseudo-liatris Cass. 
d. Basal leaves broad, linear or lanceolate. 
e. Basal leaves lanceolate, usually pubescent, not leathery, 
ascending.................... 4. C. tomentosus (Michx.) T. & G. 
e. Basal leaves linear or linear-lanceolate, glabrous, 
leathery, forming a depressed rosette (endemic to 
Konda): eoo —— . 5. C. carnosus (Small) James. 
a. Phyllaries scarcely, if at all imbricate (in 1-2(-3) series), 
glabrous or with few glandular trichomes or resinous atoms, 


the margins eciliate; involucre 4-5 mm. high....... Trilisa (Cass.) Cass. 
f. Stems glabrous............. LL. 6. T. odoratissima (Michx.) Cass. 
f. Stems pubescent................ 7. T. paniculata (Michx.) Cass. 


—DEPARTMENT OF BOTANY, UNIVERSITY OF GEORGIA. 


LITERATURE CITED 


BENTHAM, G. 1873. Genera plantarum. 2: 172. 
Cassini, H. 1816. Bulletin Société Philomathique. 198. 
— ——— —- 1818. Bulletin Société Philomathique. 140. 
1820. Dictionnaire Science Naturelle 16: 10. 
—————— 1828. Dictionnaire Science Naturelle 55: 310. 
GarsER, L. O. 1946. The genus Liatris. Ruopora 48: 165-183, 216- 
~ 263, 273-326, 331-382, 393-412. 
1949. Chromosome Studies in Liatris. I. Spicatae and 
Pyenostachyae. Amer. Journ. Bot. 36: 122-135. 
——— ——— 1950. Chromosome Studies in Liatris. II. Graminifoliae 
and Pauciflorae. Amer. Journ. Bot. 37: 414-423. 
1954. Studies in the Kuhniinae (Eupatorieae) II. Jour. 
Arnold Arb. 35: 87-133. 
McVaueu, R. 1945. The genus Triodanis Rafinesque, and its relation- 
ships to Specularia and Campanula. Wrightia 1: 13-52. 
MicHaAux, A. 1803. Flora boreali-americana 2: 93-94. 
Rosinson, B. L. 1913. A generic key to the Composite-Eupatorieae. 
Proc. Amer. Acad. Arts and Sci. 49: 429-437. 
1917. A monograph of the genus Brickellia. Mem. Gray 
Herb. 1: 1-151. 
SuiNNERS, L. H. 1943. Revision of the genus Kuhnia L. Wrightia 1: 
122-144. 


122 Rhodora [Ver. 60 


ScuuBERT, B. G. 1942. Willdenow's species plantarum and Michaux's 
flora boreali-americana: dates of publication. RHODORA 44: 147-150. 

SMALL, J. K. 1924. Plant novelties from Florida. Bull. Torr. Bot. Club 
51: 379-393. (392). 

——————- 1933. Manual of the southeastern flora. 1304. 1-1554. 

WALTER, T. 1788. Flora caroliniana. 196. 1-263. 

WinLpENOw, K. 1803. Species plantarum 3(3): 1637. 


CHROMOSOME RACES IN THE CHRYSANTHEMUM 
LEUCANTHEMUM COMPLEX! 


GERALD A. MULLIGAN? 


In 1954, as the result of chromosome studies on Canadian weeds, 
the presence of two chromosome races in North American material 
of Chrysanthemum leucanthemum L. s.l., oxeye daisy, was detected. 
Subsequent study revealed that the abundant and widespread 
oxeye daisy of North America is diploid with 18 somatic chromo- 
somes. Tetraploid plants do occur on this continent but the occur- 
rences are not widespread. 

I determined the number of chromosomes in 36 lots of material 
from different locations in Nfld., Lab., P.E.L, N.S., N.B., Que., 
Ont., B.C., and Me. A somatic number of 18 was determined on 
32 lots of this material and the other 4 lots had 36 somatic chromo- 
somes. The tetraploid plants were grown from seed collected at 
Batiscan, Lauzon and Lennoxville in the Province of Quebec and 
at Tidehead, New Brunswick. Cooper and Mahony (1935) 
counted 18 meiotic chromosomes on material from the campus 
of the University of Wisconsin and Martin and Smith (1955) 
counted 18 somatic chromosomes in material from Corvallis, 
Oregon. Three chromosome races of C. leucanthemum L. s.l., with 
somatic chromosome numbers of 18, 36 and 54, occur in Europe. 
I counted 36 chromosomes on material received from France and 
the U.S.S.R. and 54 mitotic chromosomes on two lots of material 
from Portugal. Other counts on European material were made by 
Polya (1950) on diploid plants and Negodi (1937), Ohrt in Tischler 
(1950) and Lóve and Lóve (1956) on tetraploid plants. Dowrick 
(1952) and Bócher and Larsen (1957) obtained somatie counts of 
18, 36 and 54 on European material. Three tetraploid counts 

! Contribution No. 1607 from the Botany and Plant Pathology Division, Science Service, 


Canada Department of Agriculture, Ottawa, Ontario. 
? Assistant Botanist (Weed Investigations). 


1958] Mulligan,—Chromosome Races 123 


were obtained on Japanese material by Tahara (1915), (1921) and 
Shimotomai (1937). 

The maximal inner width of pollen grains of the diploid, tetra- 
ploid and hexaploid plants counted was measured? and a correla- 
tion was found to exist between size of pollen grains and chromo- 
some numbers. The pollen grains of diploid plants ranged from 
16.3u to 19.5u, tetraploids from 19.54 to 22.8u and hexaploids from 
22.8u to 24.4u. Pollen grains from 191 Canadian and United 
States herbarium specimens were measured. None of these speci- 
mens had pollen grains in the hexaploid size range. A total of 162 
herbarium specimens collected in Nfld., Lab., N.S., P.E.I., N.B., 
Que., Ont., Man., Sask., Alta., B.C., Mass., Vt., N.Y., Va., W.Va., 
Mich., Minn., Colo., Mont., Ida., Wash., Nev., and Calif. had 
pollen grains in the diploid size range. The remaining 29 sheets 
had pollen grains that fell within the tetraploid size range. A 
total of 17 of these *'tetraploids" had been collected in the area 
between Quebec City and Gaspé or in the vicinity of Granville 
and Digby, Nova Scotia. The other 12 herbarium sheets with 
tetraploid-size pollen grains were collected from other locations 
in Lab., Que., Ont., Man., B.C., Minn., and Wash. The inner 
diameters of pollen grains from 15 European herbarium specimens 
were examined and 12 sheets had pollen grains in the tetraploid 
size range, 2 in the diploid range and 1 in the hexaploid range. 

It appears, from chromosome counts and pollen data, that most 
of the North American plants of C. leucanthemum L. s.l. are diploid 
although a small amount of our material is tetraploid. In Europe, 
the common C. leucanthemum L. s.l. is not diploid but tetraploid. 
Diploid plants seem to be slightly less common than tetraploids 
in Europe and hexaploids are rare. Dowrick in personal corre- 
spondence, dated December 2nd 1955, wrote: “Of the European 
C. leucanthemum plants which I have counted 2n = 36 is by far 
the most frequent number. The 2n = 54 plants came from Swit- 
zerland and the one count of 2n = 18 from plants obtained from 
Ireland."  Bócher and Larsen (1957) counted 26 lots of plants 
from 10 European countries and 9 lots were diploid, 16 tetraploid 
and 1 lot hexaploid. 

'The morphological differences between my diploids and tetra- 
ploids are similar to those given by Fernald (1903) when he de- 


? Pollen was removed from open disk florets and stained with cotton blue in lacto-phenol. 
Only well stained pollen showing three open pores were measured. 


124 Rhodora [Vor. 60 


scribed the characters differentiating his Chrysanthemum leucan- 
themum L. var. subpinnatifidum from what he considered the 
typical C. leucanthemum L. He recognized that the common 
ox-eye daisy of North America had in general a uniformly. different 
type of foliage from the common plant of Europe and named the 
common North American plant var. subpinnatifidum. | Fernald's 
variety has the characters of my diploid plants; the basal leaves 
generally coarsely and irregularly toothed and the middle and 
upper leaves usually narrowly oblong or oblanceolate, conspicu- 
ously subpinnatifid at the base. The characters he gave for the 
plant that is localized in North America and most common in 
Europe, the so called C. leucanthemum L., correspond to the 
morphological characters found in my tetraploids. The basal 
leaves are usually spatulate-obovate and closely and regularly 
crenate and the middle and upper leaves are usually oblong or 
oblanceolate, coarsely crenate or dentate above with larger spread- 
ing teeth at the base. 

Bócher and Larsen (1957) examined the type specimen of 
Chrysanthemum leucanthemum L. in the British Museum. This 
plant although lacking stem leaves had diploid size pollen grains 
and was considered by the authors to be morphologically similar 
to their diploid plants. They include in their paper an excellent 
photograph of a diploid plant originally collected at Edenderry, 
Eire. This photograph and their description of the European di- 
ploids convince me that their diploids are morphologically indis- 
tinguishable from my diploids and the common oxeye daisy of 
North America. It can be concluded that our common oxeye daisy 
is Chrysanthemum leucanthemum L. s. str. Bócher and Larsen 
believe that European tetraploid plants should be placed in 
Chrysanthemum ircutianum Turez. s.l. From the photographs and 
description in their paper it is evident that our North American 
tetraploids are very similar in morphology to their tetraploids. 
Unless the size of pollen grains is known, it is often impossible to 
positively identify tetraploids on anything but a complete her- 
barium specimen. Therefore, it appears premature to regard the 
tetraploids as a separate species. 

In summary, the common oxeye daisy of North America has 
a somatic chromosome number of 18 and appears to be the typical 
Chrysanthemum leucanthemum L. s. str. Plants with a somatic 
number of 36 also occur on this continent but the stands are rela- 


1958] Long,—Distribution of Ohio Compositae 125 


tively few and localized. Plants with somatic numbers of 18, 36 
and 54 occur in Europe; the tetraploid plants are somewhat com- 
moner than the diploids and hexaploids are rare. 


LITERATURE CITED 


Bocuer, Tyce W. and Kar Larsen. 1957. Cytotaxonomical studies 
in the Chrysanthemum leucanthemum complex. Watsonia 4: 11-16. 
Coorzn, D. C. anp K. L. Manony. 1935. Cytological observations on 

certain Compositae. Am. J. Botany 22: 843-848. 

Downrck, G. J. 1952. The chromosomes of Chrysanthemum, I: the 
species. Heredity 6: 365-375. 

FERNALD, M. L. 1903. Chrysanthemum leucanthemum and the Ameri- 
can white weed. Rhodora 5: 177-181. 

Love, ASKELL AND Doris Love. 1956. Cytotaxonomical conspectus 
of the Icelandic flora. Acta Horti Gotoburgensis 20: 211. 

Martin, RicHarD W. AND Frank H. SurrH.. 1955. Megagametophyte 
development in Chrysanthemum leucanthemum L. Bot. Gaz. 116: 
243-249. 

Necop1, Giorgio, 1937. Nuovi reperti cariologici su Fanerogame. 
Atti. Soc. Natur. e. Modena 68: 9-11. 

PoLva, LaszrLo. 1950. Magyarországi novénfajok kromoszómaszamai 
II. Ann. Biol. Univ. Debrecen 1: 46-56. 

SHIMOTOMAI, NAomAsa. 1937. Chromosomenzahlen bei einigen Arten 
von Chrysanthemum. Z. indukt. Abstamm. u. Vereb. 74: 30-33. 

TaHARA, Masato, 1915. Cytological studies on Chrysanthemum. 
Bot. Mag. (Tokyo) 29: 48-50 & (5)-(16). 

TaAHARA, Masato. Cytologische studien an Einigen Kompositen. J. 
Coll. Sci. Imp. Univ. Tokyo 48: 1-53. 

TISCHLER, GEoRG. 1950. Die Chromosomenzahlen der Gefasspflanzen 
Mitteleuropas, Dr. W. Junk, 's-Gravenhage. p. 152. 


NOTES ON THE DISTRIBUTION OF OHIO COMPOSITAE: 
I. HELIANTHEAE, ANTHEMIDEAE! 


Rosert W. Lona 


This series of observations was made during the preparation of 
“A Preliminary List of the Compositae of Ohio." The plants 
named below are those whose occurrence in Ohio is poorly under- 
stood judging from the information given in Gray’s Manual 
(1950) and The New Britton and Brown Illustrated Flora (1952). 
Most of these plants were introduced into Ohio as weeds either 
from Europe or from western states. This illustrates, however, the 

1 This study was aided in part by a grant from the Ohio Academy of Science. 


2 Lone, RoBERT W. (1957) A preliminary list of the Compositae of Ohio. The Ohio 
Flora Comm., Ohio State University, Columbus, Ohio. 


126 Rhodora [Vor. 60 


changing composition of the state flora, especially with respect to 
the Composilae. 

All specimens cited here are deposited in the Herbarium of the 
Ohio State University, and the identifications all have been verified 
by the writer. 

HELIANTHEAE 


Ambrosia bidentata Michx. This western species of prairies and other 
dry places is apparently known only from the extreme southern portion 
of the state, in the Allegheny Plateau. It was undoubtedly introduced 
from regions west, but the evident delimitation to the unglaciated portion 
of Ohio cannot be explained at this time. COLLECTION DATA: Jackson 
Co. Madison Twp., in a cornfield four miles NE. Thurman, Floyd Bartley 
and Lawrence Hicks, Aug. 14, 1955. Also, specimens were examined 
from Adams, Gallia, and Lawrence counties. 

Ambrosia psilostachya DC. var. coronopifolia (T. & G.) Farw. 

Another western ragweed, this plant appears to be limited to the extreme 
northern part of the state. It is no doubt adventive from the West, 
with collections being made chiefly from railroad yards, and other waste 
places. COLLECTION DATA: Lake Co., N. Y. Central R. R. at Perry, 
Fred J. Tyler, July 23, 1933. Also, Huron and Ottawa counties. 

Coreopsis verticillata L. This is a plant of dry woods and is found 
in the southeastern states. Its occurrence in Ohio is considerably north 
of its center of distribution. Only one collection was seen and the species 
is probably very rare. COLLECTION DATA: Clark Co., Silver Lake near 
New Carlisle, John H. Schaffner, Aug. 30, 1929. 

Helianthus angustifolius L. This species is known only from one 
county at the southeastern edge of the state. It is probably an escape 
from cultivation, or otherwise introduced. Typically, this plant grows 
in moist places in southeastern United States, but extends inland to 
Kentucky and Indiana. There are two separate collections from the 
same county in Ohio. COLLECTION DATA: Washington Co. in an old 
field, 3 mi. N. Marietta, Floyd Bartley, Oct. 20, 1951. 

The genus Helianthus is well-represented in the state and a number 
of putative hybrids have been identified. They are Helianthus X 
ambiguus T. & G. pro. sp., H. x luxurians Watson pro. sp., H. giganteus 
x mollis, H. occidentalis X grossesserratus, and H. petiolaris X annuus. 
Helianthus brevifolius Watson was based on a collection from Lake 
county and the type has been examined. The status of this name is 
not clear at present, but it appears to refer to a hybrid of Helianthus 
grosseserratus and H. divaricatus (H. X divariserratus Long), judging 
from the size, shape, and arrangement of leaves. COLLECTION DATA: 
Lake Co., Richmond, Otto Hacker, 1894, (TYPE). 

Rudbeckia hirta L. var. Brittonii (Small) Fern. This variety is dis- 
tinguished from the typical one by the presence of oblong phyllaries. 
Known from West Virginia, this record represents a westward extension 
for the variety. COLLECTION DATA: Coshocton Co. Harold N. Moldenke 
13298, July 25, 1942. 


1958] Long,— Distribution of Ohio Compositae 127 


Rudbeckia tenax Boynt. & Beadle. A single collection from southern 
Ohio, this would constitute an eastward range extension for the species, 
previously reported for Indiana and Illinois. COLLECTION DATA: Adams 
Co. SW. corner Oliver Twp., openings, post oak area, E. Lucy Braun. 

Silphium laciniatum L. Schaffner has noted on a collection from 
Franklin Co. that this plant is probably not native to Ohio, but rather 
is adventive from the West. In his Revised Catalogue,? however, he 
lists it as apparently indigenous. Its chief distribution occurs in the 
prairies of Michigan and North Dakota, Oklahoma, and Texas. It is 
known from widely separated areas in Ohio. One of the collections is 
from the unglaciated, southern tip of the state. COLLECTION DATA: 
Lawrence Co., collected in an old field along route 141, 1.6 mi. S. Wilgus, 
200 plants in one patch; Floyd Bartley and Lawrence Hicks, Aug. 3, 1952. 
Also, specimens from Franklin and Summit counties. 


ANTHEMIDEAE 


Achillea Ptarmica L. A single collection with “flowers not doubled,” 
probably escaped from local cultivation. In general, this collection 
is south of the chief Midwestern area of distribution for this species. 
COLLECTION DATA: Franklin Co., W. H. Camp, June 1, 1934. 

Anthemis arvensis var. arvensis L. A single collection, but at this 
location very abundant as a weed, evidently. According to the distribu- 
tion given in Gray’s Manual this would be a westward range extension. 
COLLECTION DATA: Highland Co., Hillsboro, B. & O. freight yards, Katie 
M. Roads, June 22, 1931. 

Anthemis arvensis var. agrestis (Wallr.) DC. Differing from the 
typical variety by the presence of chaff shorter than disk flower; more 
common than the preceding, but from widely scattered localities in 
Ohio. COLLECTION DATA: Greene Co., in a yard near Fairborn, Clara 
Weishaupt, June 25, 1953. Other collections from Lorain and Auglaize 
counties. 

Anthemis mixta L. The occurrence of this plant as a weed is a west- 
ward extension of the range given in the manuals; A single collection. 
COLLECTION DATA: Lake Co. Painesville, Ohio; Otto Hacker, July 28, 
1901. 

Anthemis tinctoria L. From four widely separated areas of Ohio, 
probably commener than indicated here. COLLECTION DATA: Marion 
Co., R. A. Dobbins, August 24, 1937. Other collections from Fairfield, 
Guernsey, and Leke Counties. 

Artemsia albula Woot. This species is not given in either manual 
and is definitely a plant of the western plains, specifically of ‘Texas, 
Colorado, and New Mexico. It was collected, however, outside of 
cultivation, arc ‘he specimen is typical in every way according to its 
description. COLLECTION DATA: Coshocton Co. along fence row, Mill 
Creek watersh , H. N. Moldenke 13320, July 28, 1942. 

Matricaria Chamomilla L. From the collections seen, this plant is 


3 SCHAFFNER, Jonn H. 71932) Revised Catalogue of Ohio Vascular Plants. Ohio Biol. 
urvey 5: 89-215. 


128 Rhodora [Vor. 60 


found in widely seattered portions of the state as an occasional weed. 
COLLECTION DATA: Pike Co., Jackson Twp., in an old field head of Toad 
Heaven Hollow; Floyd Bartley, May 17, 1947. Other collections from 
Greene, Lake, Lawrence, and Ottawa counties.—DEPARTMENT OF BOTANY, 
OHIO WESLEYAN UNIVERSITY, DELAWARE, OHIO 


A TAXONOMIC STUDY OF THE GENUS PHYSALIS 
IN NORTH AMERICA NORTH OF MEXICO 


U. 'T. WATERFALL 
(Continued f om p. 114) 


KEY TO GROUPS OF SPECIES OR TO UNIQUE SPECIES 


1. Corolla yellow, yellowish-green or white, with or without darker 
spots; plant surfaces without crystalline vesicles; enlarging 
ovules all of one kind. 
2. Corolla with broad shallow sinuses between the lobes; fruit- 
ing calyx reddish. ...... lll liiis 1. P. Alkekengi. 
2. Corolla truncate; fruiting calyces not red. 
3. Plants covered with stellate or variously branched tri- 
chomes, or glabrous with a few stellate hairs on the sepals 
or sometimes on the leaf margins...................... GROUP I. 
3. Plants nearly glabrous, or variously hairy, but branched 
hairs, if present, very small and inconspicuous and usually 
much less numerous than the short unbranched hairs mixed 
with them. 
4. Anthers (2) 3-5 mm. long; perennials excepting P. 
ixocarpa and P. Wrighti............. csse GROUP II. 
4. Anthers (.5) 1-2.3 (2.8) mm. long; annuals 
5. Plants nearly glabrous, usually with a few short 
curved or appressed hairs on the sepals or young parts. GROUP III. 
5. Plants long-hairy, sometimes with shorter hairs or 
glands intermixed. . 2.2... 0000... 0c cc ess GROUP IV. 
1, Corolla blue to purple (rarely white), or yellow and with plump 
seedlike corky bodies mixed with the reniform seeds. 
6. Corolla blue to purple (rarely white), rotate; no corky bodies 
mixed with the seeds; herbage with few to many crystalline 


vesicles, sometimes giving it a scurfy look............ 21. P. lobata. 
6. Corolla yellowish, funnelform; plump, rounded, corky seed- 
like bodies mixed with the reniform seeds.......... 22. P. Carpentert. 
GROUP I l 


1. Hairs stellate, each ray sometimes rebranched, the verticils 
sometimes in more than one series, and sometimes the branches 
irregularly arranged; or plant with a coat of short stellate hairs, 
plus either long-stiped branched hairs, or simple hairs. 

2. Leaves ovate to narrowly linear, their blades mostly 2.5-6 
times longer than the petiole, sometimes decurrent on it; 
corolla usually not dark spotted, or with spots not very 
prominent in herbarium specimens (rarely with prominent 
dark spots and more or less spathulate leaves); maritime 


19581. Waterfall, —Genus Physalis in N. America 129 


plants extending from southeastern Va. to the Gulf Coast 
of Texas. 
3. Leaf blades ovate, spathulate, lanceolate or linear-lanceo- 
late (sometimes linear in intergrades with P. angustifolia); 
mostly stellate-vestite, but nearly glabrous in one form of 
var. Elliottin. 
4. Leaf blades ovate to spathulate. 
5. Leaves with definite petioles usually about one-third 
to one-fourth the length of the blade; whole leaf (1.7) 
2-3 (3.4) times longer than wide; se. Va. to Fla. 
6. Leaf blades usually 2-4 (5) em. wide. 
2a. P. viscosa, var. maritima, f. maritima. 
6. Leaf blades usually 5-7 cm. wide. 
2a. P. viscosa, var. maritima, f. latifolia. 
5. Leaves tapering to the base, or extending gradually 
into winged petioles; whole leaf (2) 2.5—4 (4.7) times 
longer than wide; Gulf Coast of Texas. 
2c. P. viscosa, var. spathulaefolia. 
4. Leaf blades lanceolate to linear-lanceolate; leaves 
mostly 2.5-10 times longer than wide. 
7. Plants stellate-vestite .... 2b. P. viscosa, var. Elliottii, f. Elliottii. 
7. Plants glabrous except on the margins or the tips of 
the sepals, and sometimes on the leaf margins. 
2b. P. viscosa, var. Elliottit, f. glabra. 
3. Leaves linear, often ca. 8, but sometimes from 1-20 times 
longer than wide; plant glabrous except the tips or margins 
OL NE RODRIG v ei coast ee ee 3. P. angustifolia. 
2. Leaves mostly ovate, sometimes reniform, ovate-deltoid, or 
ovate-lanceolate, their blades mostly (1.2) 1.5-2 (3) times 
as long as their petioles; corolla dark-spotted; mostly plants 
of the south central plains, but extending to the Gulf Coast 
of Texas. 
8. Hairs 1-4 mm. long on at least the calyx or the base of the 
stem, in addition to stellate hairs covering herbage. .4. P. variovestita. 
8. Plants without hairs 1-4 mm. long intermingled with — 
shorter stellate ones. 
9. Flowering calyces (6) 7-10 mm. long; vestiture forming 
a dense mat, at least beneath the leaves; leaves dentate. 
2d. P. viscosa, var. mollis. 
9. Flowering calyces mostly (3) 5-7 (9) mm. long; vestiture 
usually sparse; leaf margins dentate, undulate or entire. 
2e. P. viscosa, var. cinerascens. 
1. Hairs jointed, those of the stem mostly 1-2 mm. long and 
spreading at right angles to the stem giving it a bristly appear- 
ance; hairs 2- or 3-branched, not having a second coat of short 
I PREFERRED cece wb esos + y's) UE Qoo de Ee. 5. P. pumila. 


GROUP II 


1. Long cord-like rhizomes present near the surface of the soil; 
found in Fla. and adjacent areas. 
2. Plants with hairs short and antrorse, sometimes viscid. 
6a. P. arenicola, var. arenicola. 
2. Plants ciliate with jointed hairs 1.5-2 mm. long. 
6b. P. arenicola, var. ciliosa. 


130 Rhodora [Vor. 60 


l. Plants with deeply buried, seldom-collected rhizomes, or 
possibly other perennating structures, or, in one species, annual. 
3. Anthers blue, ca. 3 mm. long; corolla blue- or purple-spotted; 

leaves lanceolate; perennials of s. Ariz. and adjacent Mexico. 

11. P. caudella. 
3. Anthers often yellow; if anthers blue, then plants not lanceo- 

late-leaved perennials with purple-spotted corollas native 

to s. Ariz. and adj. Mexico. 

4. Corolla usually dark-spotted near the base of its limb; 
flowering peduncles 3-15 mm. long; if corolla not notice- 
ably dark-spotted, then flowering peduncles ca. 3-8 mm. 
long. 

5. Flowering peduncles usually 10-15 mm. long; corolla 
limb usually not reflexed when fully open; plants pri- 
marily of the eastern and northern U. S. 

6. Vestiture villous; hairs jointed. 

7. Filaments as wide as the anthers to about one-third 
as wide, sometimes clavate; leaves blunt to pointed; 
anthers yellow to light blue. 

8. Anthers (3) 3.5-4.5 mm. long; filaments often 
clavate. 
9. Bases of stems not thickened and subligneous. 
10. Vestiture of stems various, but not of 
abundant hairs 2-4 mm. long. 
7a. P. heterophylla, var. heterophylla. 
10. Vestiture of abundant hairs 2-4 mm. long; 


Ala. and Fla........... 7c. P. heterophylla, var. villosa. 

9. Bases of stems thickened and subligneous; 
se. Va................. 7b. P. heterophylla, var. clavipes. 

8. Anthers (2) 2.5 (3) mm. long; filaments not 
clavate... ..........0000. 9a. P. virginiana, var. virginiana. 

7. Filaments slender; leaves acuminate; corolla spots 
and anthers deep blue-purple; introduced... ..... P. peruviana. 


6. Vestiture not villous. 
11. Flowering calyx campanulate, the lobes spreading, 
4-5 mm. wide at base and 15-20 mm. wide at tips. 
9g. P. virginiana, var. campaniforma. 
11. Flowering calyx not as above. 
12. Hairs of stem short, retrorse. 
9a. P. virginiana, var. virginiana. 
12. Hairs not short and retrorse, often antrorse. 
13. Anthers light blue, or tinged with light blue. 
14. Fruiting calyx 2.5-3.5 cm. long and 2-3 em. 
wide. .9b. P. virginiana, var. subglabrata, f. subglabrata. 
14. Fruiting calyx 4-5 cm. long and 3-4 cm. 
wide. .9b. P. virginiana, var. subglabrata, f. macrophysa. 
13. Anthers yellow. 
15. At least a few stiff spreading hairs about 1 
mm. long on the flower buds,leaf margins or 
stems; thick-leaved plants of the prairie 
region westward, usually in sand. 
9e. P. virginiana, var. hispida. 


1958] Waterfall, —Genus Physalis in N. America 131 


15. Plants without stiff spreading hairs. 
16. Principal leaves ovate; plants nearly 
glabrous usually several-branched from 
the base, and spreading; s. Tex. 
9c. P. virginiana, var. texana. 
16. Principal leaves usually lanceolate to 
linear, if ovate then plant not with sev- 
eral spreading branches from near the 
base. 
17. Plants usually single-stemmed, erect; 
larger basal leaves usually 5-10 em. 
longe s eae 9d. P. virginiana, var. sonorae. 
17. Plants usually branched from the base; 
larger basal leaves usually 4-5 cm. 
long; s. Colo. ...9f. P. virginiana, var. polyphylla. 
5. Flowering peduncles usually 3-8 mm. long; corolla limb 
often reflexed when fully open; plants primarily of the 
southern Rocky Mts. and westward. 
18. Plants with long jointed hairs mixed with shorter 
hairs, or with glandular ones, or with only long 
jointed hairs. 
19. Flowering calyx 8-11 mm. wide; anthers 1.5-3 
mm. long; leaf blades frequently rotund; mostly 
in the north central prairies extending west into 
the Rockies............... 10b. P. hederaefolia, var. comata. 
19. Flowering calyx 4-8 mm. wide; anthers 3-4 mm. 
long; sw. Tex. to Colo. and westward. 
10a. P. hederaefolia, var. hederaefolia. 
18. Plants without long jointed hairs; short hairs, or 
glandular ones present. 
20. Anthers yellow, not prominently twisted after 
dehiscence; perennials. 
21. A few short stiff branched hairs with a spread of 
ca. 1 mm. present at least on the calyces, some- 
times abundant; leaves ovate to lanceolate. 
10c. P. hederaefolia, var. cordifolia. 
21. Small branched hairs not present; leaves mostly 


ovate............ 10a. P. hederaefolia, var. hederaefolia. 
20. Anthers blue, 3 mm. long; strongly twisted after 
dehiscing; annual8............ isle esses 13. P. ixocarpa. 


4. Corolla not dark spotted, or with slightly darkened spots 
which hardly show when dry, sometimes turning blue when 
dry. 
22. Flowering peduncles 3-8 (10) mm. long, shorter than 
the flowers, or about equalling them. 
23. At least some of the hairs short, stiff and branched. 
10c. P. hederaefolia, var. cordifolia. 
23. None of the hairs short, stiff and branched. 
10a. P. hederaefolia, var. hederaefolia. 
22. Flowering peduncles (10) 12-20 (50) mm. long, some- 
what longer than the flowers to several times their 
length. 


132 Rhodora 


24. Corolla rotate, with 5 hairy pads exposed on its limb 
near the short tube; anthers blue, usually 2.5-3 mm. 


[Vor. 60 


long; annuals....... sse ne 14. P. Wrightii. 


24. Corolla not rotate with 5 hairy pads exposed on its 
limb; anthers not blue and 3 mm. long; perennials. 
25. Corolla remaining yellow when dried; leaves thick, 
often entire; flowering calyces usually 4-6 mm. 
long on peduncles from little longer than, to 6 or 7 
times their length ........ 12a. P. crassifolia, var. 
25. Corolla often drying with a blue tinge; leaves thin, 
often toothed; flowering calyces usually 3-4 mm. 
long on peduncles 5-10 times their length. 


12b. P. crassifolia, var 


GROUP III 
l. Fruiting calyx rather rounded, or 10-ribbed, but not strongly 
5-angled. 
2. Leaves ovate to ovate-lanceolate; corollas usually 6-10 mm. 
long. 
3. Flowering peduncles usually 5-15 mm. long (as much as 3 
times the length of the calyx); fruiting peduncles usually 
20-30 mm. long, shorter than, to equalling, the fruiting 
calyces which are 25-35 mm. long; flowering calyces 
usually 4-5 mm. long with teeth 2-2.5 mm. long; s. U.S. 


15a. P. angulata, var. 


3. Flowering peduncles usually 15-40 mm. long (3-13 times 
the length of the calyces); fruiting peduncles usually 20-40 
mm. long, equalling to 3 times as long as the shorter fruit- 
ing calyces which are 20-25 mm. long; flowering calyces 
usually ca. 3 (4) mm. long with teeth ca. 1 mm. long; s. 


2. Leaves lanceolate to linear-lanceolate; corollas usually 4—5 


crassifolia, 


. versicolor. 


angulata. 


Meer ehe 15b. P. angulata, var. pendula. 


mm. long; sw. U. S..... 02... .00005. 15e. P. angulata, var. lanceifolia. 


1. Fruiting calyx sharply and strongly 5-angled; corolla with 5 
evident dark spots 


GROUP IV 

1. Corolla dark spotted; anthers usually some shade of blue or 
purple. 
2. Fruiting calyces 1.5-2.5 (3) cm. wide. 

3. Anthers (1.2) 1.5-2 mm. long; plants without capitate- 
glandular hairs, but sometimes viscid-glandular or with 
sessile glands. 

4. Leaves having a greyish surface, often with “mealy” or 
sessile glands; leaf blades usually toothed nearly to the 


MMC 16b. P. pubescens, var. glabra. 


base; mostly northeastern U. &...16d. P. pubescens, var. grisea. 


4. Leaves not greyish, not having sessile glands. 
5. Leaves usually toothed nearly to the base with 5-8 
teeth on each side of the seldom translucent blade; 


widespread..............-- 16a. P. pubescens, var. pubescens. 


5. Leaves with few teeth, 3-4 on each side, or entire; 
blades mostly flaccid and translucent. 


16a. P. pubescens, var. integrifolia. 


1958] Waterfall, —Genus Physalis in N. America 133 


3. Anthers (.3) 1-1.5 mm. long; capitate-glandular hairs 
usually mixed with long jointed ones.17. P. foetens, var. neomexicana. 
2. Fruiting calyces (2.5) 3-4 em. wide; s. Ariz............. 18. P. latiphysa. 
1. Corolla yellow, unspotted, or sometimes slightly dark-tinged. 
6. Flowering peduncles 2-5 mm. long, about equalling to twice 


aslongasthecalyces..................Lue lesus. 19. P. missouriensis. 
6. Flowering peduncles mostly 15-20 (30) mm. long, 4-5 (7) 
times the length of the calyces.................000005. 20. P. Greenei. 


l. Physalis Alkekengi L., Species Plantarum 183. 1735; incl. P. 
Francheti Mast. in Gard. Chron. 2: 434 and 441. 1894. 


Plants perennial, erect, usually unbranched, 30-60 cm. tall, glabrous or 
with a few scattered long hairs; leaves broadly ovate, or ovate-rhombie, the 
blades usually 5-12 em. long and 4-9 em. wide on petioles 2-4 em. long; mar- 
gins of the leaf blades from entire to irregularly few-toothed or undulate- 
dentate; flowers white, slightly 5-lobed with broad shallow sinuses about 2 
mm. deep between the apices of the lobes; corollas 10-15 mm. long and 15-25 
mm. wide; anthers 2.5-3 mm. long on slender filaments; flowering calyces 4—7 
mm. long and 4-5 mm. wide, densely hairy with jointed trichomes about 1-1.5 
mm. long; fruiting calyces reddish, 3-5 cm. long and 2.5-4 em. wide, pendent 
on peduncles 2-3 em. long. 


TYPE: Not seen; Linnaeus states, ‘‘Habitat in Italia.” 

This species is cultivated or escaped, in northeastern U. S.; flowering in 
June; 19 sheets examined. 

2. Physalis viscosa L., Species Plantarum 183. 1753; other synonymy 
under the varieties to which the names are referred. 


Perennials covered with stellate, or several-branched hairs, or nearly glabrous 
and having stellate hairs only on the calyces; leaves from ovate to linear- 
lanceolate, petiolate or with blade tapering to the stem; corolla yellowish, 
funnelform, with or without darker spots on the limb near its Ease, 8-20 mm. 
long; anthers yellow, about 3 mm. long; flowering calyx 3-10 mm. long on 
peduncles 10-20 mm. long; fruiting calyx 2-5 em. long and 1.5-4 em. wide on 
peduncles 1-4 cm. long. 


P. viscosa L., ssp. viscosa. P. viscosa L., loc. cit., as limited to the 
South American plants. Linnaeus cited P. viscosa from ‘Virginia, 
Bonaria." Since (1) only one element of this species, the part described 
by Chapman as P. maritima, barely extends into southeastern Virginia, 
since (2) it hardly matches the photographs of the Linnean types, and 
since (3) previous authors have restrieted the application of the specific 
name, in its strictest sense, to the plants of South America, the present 
author believes that it is best to so delimit it. 

South American material is usually more sparsely vestite with finer 
stellate hairs than most of ours, and has corollas slightly spotted to 
unspotted. 

Occasionally in our populations of P. viscosa, sens. lat., there appears a 
specimen which can hardly be distinguished from South American collec- 
tions. Whether these are introductions from that continent, or represent 
gene combinations from within our population which produce phenotypes 


134 Rhodora [Vor. 60 


similar to the South American plants, it is impossible for the author to 
determine. 


Some examples are: ALABAMA: Alabama Co.: Mohr 26 (Ny); Texas: Brazoria 
Co.: Young Apr. 22, 1918 (rex); Houston Co.: Fisher July 25, 1914 (uc); 
Walker Co.: Warner 29 (ny). 


P. fuscomaculata de Rouville ex Dunal, at least as to the few collections 
seen, is included here. 

P. viscosa ssp. maritima (M. A. Curtis) Waterfall, comb. et stat. nov., 
based on P. maritima M. A. Curtis, Am. Journ. Sci. ser 2. 1: 407. 1849. 
Under this subspecies are included the maritime varieties listed below. 

2a. P. viscosa var. maritima; P. Walteri Nuttall, Journ. Acad. Nat. 
Sci. Phila. 7: 112. 1834; type from “South Carolina" (PH); P. maritima 
M. A. Curtis, loc. cit.; P. viscosa L., var. maritima (Curtis) Rydberg, 
Mem. Torr. Bot. Club 4: 357. 1896. 

The varietal name is used above as a tautonym of ssp. maritima. How- 
ever, since the author is utilizing the varietal concept for taxa with both 
morphological and distributiona! differences, taxa which he considers the 
principal subdivisions of complex species, and is using the subspecific 
catagory as an aggregation of similar varieties, he would prefer to transfer 
the specifie name, in this case maritima, to the varietal status, letting the 
tautonym, without author citation, belong to the aggregate subspecies. 
This does not seem permissible under Article 15 of the International Rules. 

Leaves ovate to spathulate, especially the upper ones; whole leaf (1.7) 
2-3 (3.4) times longer than wide; from sparsely to usually densely vestite. 

TYPE: M. A. Curtis, seacoast, North Carolina (au); probable isotype, 
"sandy seacoasts" (NY). 

This taxon grows on sandy seacoasts, southeastern Va. to Fla.; usually 
flowering in March, April and May, but sometimes in winter in the south- 
ern part of its range; 120 sheets of 93 collections seen. 

Occasional is a large, broad-leaved form with leaf blades about 5-7 cm. 
wide as contrasted to the usual width of 2-4 (5) em. in forma maritima 
(supra). It may be described as P. viscosa f. latifolia Waterfall, f. nov., 
laminis 5-7 em. latis. The type is Small, Mosier and DeWinkeler 10892 
(ny); isotype (GH); north part of Jupiter Island, Florida. Also seen from 
Florida were: Dade Co.: Moldenke 384 (puke), Small 2116 (ny). 

2b. P. viscosa var. Elliottii (Kunze) Waterfall, comb. et stat. nov., 
based on P. Elliottii Kunze, Linnaea 20: 33. 1847. 


Leaf blades lanceolate to linear-larceolate; leaves mostly 2.5-10 times 
longer than wide; plants covered with stellate Fairs, or rearly glal rous; corol- 
las yellow, usually not dark-spotted, but scmetimes prominently so. 


TYPE: Rugel June 1843, “Ad ostium fluvii St. Marks in Florida” (NY). 
The type belongs to the vestite phase although it is only sparingly stellate 
on the leaf-surfaces and the stem. 

This taxon grows in sands, Florida; apparently flowering throughout the 
year. 

P. viscosa f. Elliottii. The following citations selected from 73 sheets of 


1958] Waterfall,-—Genus Physalis in N. America 135 


58 collections: FLoripa: Broward Co.: Moldenke 479 (DUKE, NY); Clay Co.: 
Williamson Aug. 1893 (pH); Dade Co.: Small and Mosier 5944 (DUKE, GH, NY); 
Moldenke 715 (ny); Hernando Co.: McFarlin 6079 (mica); Highlands Co.: 
Correll and McFarlin 6219 (DUKE); Hillsboro Co.: Churchill Mar. 21, 1923 (Gu, 
PH); Indian River Co.: Small 8894 (Ny); Lee Co.: Mosier July 1928 (DUKE, NY); 
Manatee Co.: Tracy 7577 (aH, Ny); Monroe Co.: Sargent 6417 (Ariz); Pinellas 
Co.: McFarlin 3653 (mica); St. John Co.: Meredith Feb. 4, 1899 (pH); Sarasota 
Co.: McFarlin July 1931 (micu). 


P. viscosa var. Elliottii. f. glabra Waterfall, f. nov., foliis glabris, 
sepalis stellato-vestitis. Stellate hairs are found on the flowering sepals, 
or at least on their margins; a few are rarely present on the margins of the 
leaves. The Type is Tracy 7608, Sanibel Island, Lee County, Florida 
(ny), isotype (GH). 

Selected from 42 sheets of 33 collections: FLoRiDA: Collier Co.: Deam 60785 
(DUKE); Dade Co.: Small 7410 (Ny); Hillsborough Co.: McFarlin 5525 (mican); 
Lee Co.: Hitchcock 237 (an, Ny); Monroe Co.: Palmer 376 (an); Pinellas Co.: 
Deam 1948 (uc); Sarasota Co.: McFarlin 6091 (mica). 


An unusual, compacted short-leaved phase is sometimes found. It is 
represented by the following: Small, Britton and DeWinkeler 2328, pine- 
land-prairie, Tamiami trail west of Miami, Dade Co., Dec. 19, 1919; Small 
8894, pinelands near Felsmere, Indian River Co., May 17, 1918; Small and 
DeWinkler 9979, Ancient sand dunes near Kuhiman, April 25, 1921, all in 
Florida. If there proves to be a population in this area, similar to the 
cited collections, it will probably justify nomenclatural recognition. 

2c. P. viscosa var. spathulaefolia (Torr.) Gray, Proc. Amer. Acad. 
Arts and Sci. 10: 67. 1875. P. lanceolata Michx., var. spathulaefolia 
Torr., Bot. Mex. Bound. 153. 1859. 


Leaf blades ovate to lanceolate to spathulate, tapering at base, or extending 
gradually into winged petioles; whole leaf (2) 2.5—4 (4.7) times longer than 
wide; corolla varying from apparently unspotted to having prominent dark 
spots; Gulf Coast of Texas; flowering irregularly throughout the year. 

TYPE: Schott 30, seabeaches, Rio Bravo (Rio Grande?), Texas (Ny). 

The following selected from 65 sheets of 54 collections: LourstANA: Calcasieu 
Parish: Palmer 7707 (pH); Texas: Aransas Co.: Tharp 1620 (oKLA, TEX); Austin 
Co.: Pennell 10271 (Ny, PH); Brazoria Co.: Cory 51057 (au); Cameron Co.: 
Clover 1724 (mich); Galveston Co.: Nelson Mar. 20, 1942 (rex); Harris Co.: 
Fisher July 25, 1914 (ny); Jefferson Co.: McVaugh 6880 (mican); Kenedy Co.: 
Tharp 48333 (OKLA, TEX); Matagorda Co.: Wright (au); Nueces Co.: Tharp, 
Johnson and Webster Dec. 3, 1948 (TEx); San Patricio Co.: Cory 51249 (au, 
us); Victoria Co.: Tharp 2512 (TEx, vc). 

The following collections have prominently dark-spotted corollas: TExas: 
Austin Co.: Tharp Apr. 8, 1939 (TEx); Brazoria Co.: Celerier 51-41 (OKLA); 
Cameron Co.: Lundell 1073 (TEx); Kenedy Co.: Lundell 8714 (GH, MICH, NY, 
uc); Willacy Co.: Johnston 54169 (TEX). 


P. viscosa ssp. mollis (Nuttall) Waterfall, comb. et stat. nov., based on 
P. mollis Nutt., Trans. Am. Phil. Soc. 5 (n.s.) 194. 1837. This is an 
inland population, here divided into two varieties, as compared with ssp. 
maritima, a maritime population divided above into three varieties. 


136 Rhodora [Vor. 60 


2d. P. viscosa var. mollis. The varietal name is used here as a tauto- 
nym of ssp. mollis, hence it is listed without author-citation. However, 
the present author believes that the varletal category is best used as the 
principal division of a species, and that the subspecifie category is most 
significantly used as a means of grouping varieties, just as a section may 
be utilized to group species within a genus. 

TYPE: not seen; isotypes: Nuttall, Arkansas (Ny, PH). 

Leaves densely stellate-tomentose, at least beneath, dentate; plant 
usually erect; flowering calyces (6) 7-10 mm. long. 

Growing in open woods, sandy areas and disturbed sites, western Arkan- 
sas, eastern Oklahoma and eastern Texas. There are many intermediates 
with var. cinerascens. Usually flowering in May and June, but speci- 
mens in flower have been collected in July and August. 


Selected from 43 sheets of 34 collections; Arkansas: Franklin Co.: Pennell 
10621 (xv, PH); Pulaski Co.: Merrill 1861 (UARK, OKLA): Sebastian Co.: 
Bigelow in 1853-54; Louisiana: Bossier Parish: Correll 10058 (GH, DUKE, NY, PH); 
Jefferson Davis Parish: Palmer 7629 (pu); Oklahoma: Choctaw Co.: Houghton 
4037 (Gu, NY); Comanche Co.: Clements 11767 (Gu); Love Co.: Hopkins 3429 
(oKL); Marshall Co.: Basler Aug. 5, 1950 (oKL); Muskogee Co.: Little 1515 
(oKL); Texas: Callahan Co.: Palmer 13811 (wis); Denton Co.: Whitehouse 
15780 (mcu); Grayson Co.: Gentry 51-392 (oxra); Tarrant Co.: Ruth 902 
(wis); Wilson Co.: Rogers, Albers and Webster 6849 (TEX). 

2e. P. viscosa var. cinerascens (Dunal) Waterfall, comb. nov., based 
on P. pensylvanica L. var. cinerascens Dunal, in De Candolle, Prodromus 
13(1): 485. 1852; P. mollis Nutt., var. cinerascens (Dunal) Gray, Proc. 
Amer. Acad. Arts and Sci. 10: 66. 1875; incl. P. mollis Nutt., var. parvi- 
folia Rydb., Mem. Torr. Bot. Club 4: 355. 1896. 


Leaves dentate to entire, varying in size, probably due, at least in part, to 
seasonal heteromorphy; plants erect to spreading, or nearly procumbent; more 
or less densely covered with stellate hairs, but rot tomentcse; flowering calyx 
(3) 5-7 (9) mm. long; small-leaved forms have been segregated as var. parvi- 
folia. 


TYPE: Dunal cited “n. 83 et 2316 Berland. pl. exs. Mex." when he 
described var. cinerascens. Since he indicated no holotype, Berlandier 
2316, cirea Matamoros urbem, April 1831 (an) is designated as Lectotype. 

'This taxon grows on prairies, plains and in disturbed habitats, primarily 
in Oklahoma and Texas, extending into Mexico; often flowering in May 
and June, but to some extent at any time during the growing season, which 
may be most of the year in southern Texas. 

Four hundred twenty seven sheets of 365 collections of var. cincerascens 
(sens. lat.) have been examined. In addition to numerous Oklahoma and 
Texas collections, the following have been seen: ARKANSAS: Fulton Co.: 
Bush 2518 (au); Kansas: Barber Co.: Rydberg and Imler 640 (NY); New 
Mexico: Chaves Co.: Earle 283 (wy); Lea Co.: Waterfall 7836 (an, OKL). 

P. pensylvanica L., Species Plantarum, ed 2, 1670, 1762, is not accounted 
for in the above synonymy. A tracing from the Linnean herbarium on a 
Canby sheet of P. viscosa in the Gray Herbarium bears the annotation 


1958] Waterfall,—Genus Physalis in N. America 137 


“pubescence very short stellular—same as in P. viscosa Hb. Linn." No 
stellate species is known from Pennsylvania. No disposition of it can be 
made from the photograph of the species from the Linnean Herbarium, 
other than to say that if it is indeed from North America, it might be a 
small, rotund-leafed phase of P. viscosa, var. maritima. 

3. Physalis angustifolia Nuttall, Journ. Acad.Nat. Sci. Phila. 7: 113. 
1834. 

Perennial from a thick woody taproot, often with many branches; leaves 
linear, (8) 10-20 times longer than wide; plants glabrous except on the 
tips or margins of the sepals; corolla yellow, unspotted to apparently 
spotted, but only lightly so, 8-20 mm. long; flowering calyx 5-10 mm. long, 
on peduncles 10-20 mm. long; fruiting calyx 2-3 em. long on peduncles 
1-3 em. long. 

TYPE: Not seen; Isotype: N. A. Ware “west Florida" (Pn). Its larger 
leaves are 6-8 em. long and about 2-3 mm. wide. 

P. angustifolia grows in coastal sands, coral soil, or pine woods, Alabama 
to Mississippi with one collection from Louisiana; it usually flowers from 
May to August, but a number of collections, particularly from Florida 
have been taken in flower in December, January and February. 

Selected from 87 sheets of 65 collections: ALABAMA: Baldwin Co.: Mohr 
March 20, 1883 (aH, UARK); Mobile Co.: Mohr 1878 (ny, pH); FLORIDA: 
Bay Co.: Banker 3670, 3679 (ny); Collier Co.: Moldenke 1006 (DUKE, NY); 
Dade Co.: Moldenke 852 (puxe, Ny); Gulf Co.: Correll and Oosting 5630 
(DUKE); Monroe Co.: Curtiss 114 (GH, MICH, NY, PH, UARK) ; Oskaloosa Co.: 
Menzel and Menzel 55-3 (wis); Santa Rosa Co.: Fassett 21141 (wis); 
Wakulla Co.: Griscom 21478 (au); Louisiana: “seashore”: Carpenter, 
July (rn); Mississippi: Jackson Co.: Baker July 25, 1897 (wy); Harrison 
Co.: Demaree 21911 (OKL, OKLA, TEX); Tracy May 8, 1898 (Ny, MIcH). 

4. Physalis variovestita Waterfall, sp. nov. Planta bivestita, pilis 
elongatis, 1-4 mm. longis, articulatis, simplicibus vel furcatis, et brevo- 
stellatis; caulibus erectis; folis petiolatis; laminis ovatis dentatis vel 
subsinuato-dentatis; ; pendunculis petiolis longioribus; corollis luteis, fundo- 
maculatis; antheris luteis ca. 3 mm. longis. 


The presence of abundant jointed hairs, 1-4 mm. long, in addition to a 
covering of stellate hairs, is the most obvious characteristic of this species. 
The long hairs are sometimes branched, sometimes simple. The plant is a 
perennial, 12-25 cm. tall, from a rhizome. The leaf blades are ovate, dentate 
to more or less sinuate-dentate, 2.5-3.5 cm. long and 2-3 em. wide on petioles 
1.5-2.3 cm. long. The nodding flowers are on peduncles 2-3 em. long. "The 
corolla is 1.5-2 cm. long and 2-3 cm. wide, with large dark markings on its 
limb near its junction with the tube. The flowering calyx is about 1 em. long 
divided about one-third to one-half of the way into ovate-lanceolate, or 
lanceolate lobes. The anthers are yellow, ovate to ovate-oblong, about 3 
mm. long. 


TYPE: Hula Whitehouse 18179, back of Rockport Tourist cottages in 
sandy soil, live-oak belt, Rockport, Aransas Co., Texas, April 21, 1947 
(mic). 


138 Rhodora [Vor. 60 


A number of collections radiating northward from this area have long 
articulated hairs present to a greater or lesser extent. These are found in 
specimens resembling both var. mollis and var. cinerascens, with a degree 
of variability in leaf size, margins, and stellate vestiture similar to that 
found in these two taxa. 

Sheets approaching the type in vestiture are: Kenedy Co.: Cory 28408 
(GH); Medina Co.: Johnston, Tharp and Turner 3401 (OKLA, TEX). 

Colleetions more widely diverging from P. variovestita, but with several 
to few long jointed trichomes present in addition to the short stellate hairs 
are: Austin Co.: Pennell 10300 (Ny, vu); Bexar Co.: Metz 477 (uc); Cald- 
well Co.: Coll. unknown (J. B. MeB.) 1931 (tex); Cameron Co.: Tharp 
1206 (tex); DeWitt Co.: Reidel Apr. 5, 1942 (GH, OKLA); Gillespie Co.: 
Bray 293 (tex); Gonzales Co.: Bogusch 1868 (TEx); Cory 8366 (au); 
Turner 8706 (vex); Jim Hogg Co.: Tharp June 17, 1928 (rex); Hidalgo 
Co.: Cameron 269 (TEx); Kenedy Co.: Johnston 58256.19 (TEx); Lundell 
8715 (au); Upshur Co.: Reverchon 3237 (wy); Victoria Co.: Coll. unknown 
Mar. 29, 1930 (rex); Waller Co.: Hall 500 (au, Ny); Wilson Co.: Cory ?795 
(au); Parks 29580 (au); Wood Co.: McMullen June 10, 1927 (TEx). 

It is postulated that a population such as described above, and exem- 
plified by the type collection, must exist in the area indicated in southern 
Texas, and that gene interchange has diluted its characteristics with those 
of P. viscosa var. mollis and var. cinerascens in an area radiating northward. 
Probably Edgar Anderson's method of extrapolated correlates (1949) 
could have been used to predict the occurrence of P. variovestita on the 
basis of the intergrades found in approaching the area in which it grows. 

5. Physalis pumila Nuttall, Trans. Am. Phil. Soc. 5 (n.s.); 193, 1836; 
P. lanceolata Michx., var. hirta Gray, Proc. Amer. Acad. Arts and Sci. 10: 
68. 18795. 

Plants perennial, 15-45 cm. tall, often branched, usually covered with 
jointed hairs 1-2 mm. long, some of which are 1- to rarely 3-branched, and 
which spread at right angles from the stem. Leaf blades ovate to ovate- 
lanceolate, or rarely lanceolate, sometimes somewhat rhombic, tapering to a 
more or less winged petiole; larger blades (4) 6-9 cm. long and (2.5) 3-5 cm. 
broad, on petioles 1-3 cm. long; leaf margins usually entire, but sometimes 
lightly and irregularly sinuate-or repand-dentate; corolla 12-20 mm. long, and 
about 15-25 mm. wide at the top; anthers usually 2.5-3 mm. long, yellow; 
flowering calyx usually 10-15 mm. long with free lanceolate-deltoid sepal tips 
about one-third as long; flowering peduncles 15-30 mm. long; fruiting calyx 
usually 15-20 mm. wide and 3-4 cm. long, much inflated around the fruit, on 
reflexed peduncles 25-40 mm. long. 

TYPE: Not seen; Isotype: Nuttall, Arkansas (pH). The isotype is repre- 
sentative of the extreme having few branched hairs. It was collected 
near the eastern limit of its range in this area. 

P. pumila grows in prairies, open woods and disturbed habitats, pri- 
marily in western Missouri, eastern Kansas, eastern Oklahoma and adja- 
cent Texas; it usually flowers in May, June, July and August, perhaps 
earlier in the southern part of its range, as fruiting specimens have been 
collected in May in Texas. 


1958] Waterfall,—Genus Physalis in N. America 139 


The 169 sheets of 146 collections examined include: ARKANSAS: Sebastian 
Co.: Armstrong 186 (TEX, UARK); Carrol Co.: Moore and litis 204 (wis); 
Washington Co.: Hill 23 (UARK); Iuuinots: Peoria Co.: Chase 3570 (NY, vc). 


6. Physalis arenicola Kearney, Bull. Torr. Bot. Club 21: 485. 1894. 


Plants perennial from cord-like rhizomes which are near the surface, usually 
15-30 em. tall, simple or branched; hairs short and antrorse, sometimes viscid, 
in var. ciliosa 1-2 mm. long, jointed, spreading and more or less abundant; 
leaf blades ovate to ovate-rhombic, the larger ones usually 2-6 em. long and 
2-4 cm. wide on petioles 1-3 cm. long; leaf margins irregularly dentate to 
sinuate or entire; corollas 10-20 mm. long, yellow with slightly darker spots on 
the limb near its base; flowering calyx 7-11 mm. long, its lobes 2-4 mm. long; 
flowering peduncle 10-25 mm. long; fruiting calyx 20-30 mm. long and 15-25 
mm. wide, much inflated around the fruit. 


TYPE: Kearney cited several collections of Nash's when he deseribed 
P. arenicola. Since no holotype was designated, the author selects the 
following from among the cited collections: George V. Nash 1170, dry sandy 
soil, high pine land, vicinity of Eustis, Lake County, Florida, July 1-15, 
1895 as the lectotype (GH); Isolectotypes: (Ny, UC). 

This species grows on sand dunes, ridges, sandy oak woods, pine woods 
and disturbed sandy areas, primarily in Florida, but also in adjacent 
Georgia and Mississippi; flowering March through August. 

6a. P. arenicola var. arenicola. Selected from 28 sheets of collections: 
Fiorwa: Alachua Co.: Wiegand and Manning 2810 (au); Brevard Co.: 
Curtiss 5713 (an, uc); Duval Co.: Curtiss 6644 (au, NY, vc); Lake Co.: 
Nash 1170 (au, vc); Levy Co.: Garber Nov. 1877 (aH); Marion Co.: 
Moldenke 1090 (pukr); Palm Beach Co.: Small 8514 (DUKE, GH); Sumter 
Co.: Curtiss 6634 (aH); Volusia Co.: Small 8692 (puKr, GH); Georgia: 
Lowndes Co.: Harper 1594 (Gu, Ny). 

6b. P. arenicola Kearney, var. ciliosa (Rydb.) Waterfall, comb. et stat. 
nov., based on P. ciliosa Rydb., Mem. Torr. Bot. Club 4: 346. 1898. 

TYPE: In describing P. ciliosa, Rydberg stated “Chapman (in Herb. J. 
Donnell 8mith, Harvard University, Columbia College, and A. W. Chap- 
man, type)." As Lectotype the author chooses a sheet (GH) showing both 
flowering and fruiting plants. Isolectotypes are: GH, a second sheet, NY 
and OKL. 


Selected from 35 sheets of 33 collections: FLoripa: Alachua Co.: Walker 
1917 (oKrLA); Brevard Co.: Small and DeWinkeler 2468 (ny); Gadsden Co.: 
Berg (ny); Hendry Co.: Moldenke 1018 (puke, Ny); Highlands Co.: Small, 
Mosier and DeWinkeler 10906 (Ny); Lee Co.: Moldenke 946 (Duke, Ny); Levy 
Co.: Oosting 139 (DUKE); Osceola Co.: Singletary Apr. 28, 1938 (DUKE); Polk 
Co.: McFarlane 5021 (micu); Santa Rosa Co.: McFarlane and Goertz June 17, 
1905 (DUKE); Sarasota Co.: Rusby April 1935 (Ny); Sumter Co.: Curtiss 6634 
(uc); Grorata: Calhoun Co.: Thorne 3321 (au); Chatham Co.: Gay (an); 
Charlton Co.: Small June 12-15, 1895; Mississippi: Jackson Co.: Skehan 
May 10, 1895 (aH). 


The following specimens, all from Florida, seem to be intermediate 
between var. arenicola and var. ciliosa: Collier Co.: Small 10477 (NY); 


140 Rhodora [Vor. 60 


Dade Co.: Small and Small 6825 (au, ny); Volusia Co.: Small 8692 (au, 
DUKE). 

7. Physalis heterophylla Nees, Linnaea 6: 463. 1831; synonymy cited 
under the varieties. 

Stems usually erect from a deeply buried rhizome, 15-90 cm. tall, simple or 
branched; herbage densely to sparsely covered with varying proportions of 
short usually viscid hairs and glandular hairs, together with long jointed hairs 
which are usually 1-2 mm. long; sometimes only a few long hairs are present; 
rarely, as in var. villosa, the stems are villous with long multicellular hairs; 
leaf blades usually broadly to narrowly ovate, or ovate-rhombic, the principal 
ones usually 5-10 cm. long and 3.5 to 6 cm. wide on petioles 3-6 em. long; 
corollas 10-18 mm. long, yellow with brownish, sordid or blue-tinged spots on 
the limb near its base; flowering calyx 7-12 mm. long, its lobes 3-5 mm. long, 
lanceolate-triangular, sometimes acuminate; anthers usually 3-4.5 mm. long, 
vellow, sometimes tinged with blue; filaments thickened, often as wide as the 
anthers, frequently clavate; fruiting calyx usually 2.5-3 em. long and 2-3 em. 
wide, much inflated around the fruit, borne on peduncles 1.5-4 cm. long. 

7a. P. heterophylla Nees, var. heterophylla P. virginiana Mill., var. 
ambigua Gray, Proc. Amer. Acad. Arts and Sciences 10: 65. 1875; P. 
nyctaginea Dunal, DeCandolle, Prodromus 13(1): 440-441. 1852; P. 
ambigua (Gray) Britton, Mem. Torr. Bot. Club 5: 287. 1894; P. hetero- 
phylla, var. umbrosa Rydberg, Contr. U. S. Natl. Herb. 3: 172. 1895; 
P. heterophylla, var. ambigua (Gray) Rydberg, Mem. Torr. Bot. Club 4: 
349. 1896; P. sinuata Rydb., in Small’s Flora: 986. 1913. 

This is an extremely variable assemblage as indicated in the preceding 
description, which, with stated exceptions, covers var. heterophylla, only 
two other localized varieties being recognized. Forms (the species or 
varieties of earlier authors) might be distinguished on the basis of denta- 
tion of leaves, or of vestiture, but many specimens would be assignable 
only on an arbitrary basis, even if some of the extremes seem quite striking. 
An example is the densely stiff-haired form often found on sands at various 
localities in the range of the species. 

TYPE: Not seen; “In collibus argillosis Pennsylvanicae Poeppig legit.” 

Habitat, range and flowering time: Open woods, prairies, hillsides, fields 
and other disturbed habitats, principally in the eastern United States and 
adjacent Canada, the prairie and plain region westward into the central 
and northern Rockies and the Great Basin; flowering from June to August 
in Canada and from April to September in Texas. 

In examining 730 sheets of 664 collections, material has been seen from: 
Ontario and Quebec, Canada and from Alabama (Jackson and Tuscaloosa 
Cos.), Georgia (Chatham and Clark Cos.), Idaho (Allen 1873), Illinois, 
Indiana, Massachusetts, Michigan, Minnesota, Mississippi (Harrison 
Co.), Missouri, Nebraska, New Hampshire, New Jersey, New York, North 
Carolina, North Dakota (Richland Co.), North Carolina, Ohio, Oklahoma, 
Pennsylvania, Rhode Island, South Carolina (Oconee Co.) South Dakota, 
Tennessee (Davidson, Frank and Rutherford Cos.), Texas, Utah (Salt 
Lake Co.), Vermont, Virginia, Washington, D. C., West Virginia, Wis- 
consin, Wyoming (Big Horn and Crook Cos.). 


1958] Waterfall,—Genus Physalis in N. America 141 


7b. P. heterophylla Nees, var. clavipes Fernald, RHODORA 49: 178. 
1947. 

TYPE: Fernald, Long and Clement 15347, sandy woods near Darden's 
Pond, northeast of Courtland, Southampton Co., Virginia, (an); isotype 
(xy). Known only from the type collection. 

7c. P. heterophylla Nees, var. villosa Waterfall, var. nov., caulibus 
dense articulato-villosis, pilis 2-4 mm. longis. 

The abundant, soft, long, jointed hairs, 2-4 mm. long, characterize this 
variety. The leaf size and margin vary in a manner comparable to var. 
heterophylla. 

TYPE: Earle June 2, 1901, moist hillsides thick woods, Lee County, 
Alabama (NY). 


Collections examined: ALABAMA: Lee Co.: Earle June 2, 1901 (xv); F. S. 
Earle May 10, 1896, Auburn (xv); FLorrpa: Gadsden Co.: Berg Summer (Ny); 
Walton Co.: A. H. Curtiss June 1886, De Funiak Springs (Ny); County unde- 
termined: W. E. Buswell April 9, 1931, Pine Woods, East Fort Meyers (Ny); 
Texas: doubtfully referred here is Tharp April 19, 1930, East Texas coast 
(TEX). 

P. heterophylla appears to intergrade with P. virginiana 1n some areas, 
producing individuals with varying indument and leaf-shape, including 
lanceolate. Such specimens are found in South Carolina from which 
Michaux described his P. lanceolata, Flora Boreali-Americana 149. 1803. 
Examples are: Gibbes, in 1834, Columbia, S. Car. (Ny); Gibbes Aug. 1835, 
South Carolina (Ny); Ravenel, Aiken, South Carolina (Ny). The photo- 
graph of the type of P. lanceolata in the Gray Herbarium appears to match 
these specimens fairly well. The author believes that it was on such a 
specimen that Michaux based his species. This leaves the population of 
the western prairies and plains, which has been passing under the name 
P. lanceolata, without a name. It will be treated under P. virginiana. 


Other collections believed to be P. heterophylla intergrades are: CONNECTI- 
cut: Bishop Sept. 1902, Norwich (an); Grorata: Harper 93, Dry fields, Clarke 
Co., June 29, 1900 (xv); Coll. unknown (herb. Schw. sub nom. “P. obscura 
Baldw., Georgia") (pH); NortH CanoriNA: Williamson Aug. 1900, Wilmington 
(PH); Small, July 1896, Summit of Paris Mt. (ny). 


8. Physalis peruviana L., Species Plantarum, ed. 2, 1670. 1762. P. 
peruviana, var. latifolia (Lam.) Dunal, in DeCandolle, Prodromus 13(1): 
440. 1852, based on P. latifolia Lamarck, Tableau Encyclopédique et 
Méthodique ... Bot. 2: 29. 1793, is the only synonymy that has been 
applied in the area under consideration. 


An erect branching perennial, densely villous but not glandular; leaf blades 
ovate, extending into an acuminate tip; corolla blue-spotted; anthers about 3 
mm. long, blue, on slender filaments. "This species resembles P. heterophyila, 
but may be distinguished by the narrow filaments and the rather strongly 
acuminate leaves, as well as by the blue anthers (sometimes the anthers are 
violet-tinged in P. heterophylla) and by the darker, bluish spots of the corolla. 


TYPE: Not seen; Linnaeus says "Habitat Limae”’. 


142 Rhodora [Vor. 60 


This species is sometimes introduced, and may rarely escape. Some 
examples are: Kidder Oct. 3, 1926, Norfolk Co., Mass. (NEBC) ; Martindale 
Sept. 1879 Camden, New Jersey (Nv); Brinkley 222, Sevier Co., Arkansas 
(rex) ?; Earle June 26, 1899, Lawrence Co., Alabama (Ny) ?. 

9. Physalis virginiana Miller, Gardener's Dictionary, ed. 8: No. 4. 
1768. The synonymy is given under the varieties. 

Stems from a deep rhizome, simple or branched; plants nearly glabrous, or 
with long hairs, or short curved trichomes; leaf blades from ovate to linear- 
lanceolate; corolla from 15-25 mm. long, yellow, dark-spotted; anthers 2-4 
mm. long, yellow or blue- or violet-tinged; filaments from one-third as wide to 
nearly equalling the width of the anthers; calyx from one-half to two-thirds as 
long as the corolla; flowering peduncles about equalling the flower to 14% 
times its length; fruiting calyx inflated, usually 25-35 mm. long and ovate to 
ovate-oblong, but sometimes much larger, particularly in one forma. 


The varieties described below seem to intergrade more or less with each 
other, making the disposition of individual specimens sometimes difficult. 
However they seem to represent natural populations, in some instances 
covering large geographic areas, which are fairly distinct as groups. 

9a. P. virginiana Miller, var. virginiana. P. virginiana Mill., var. 
intermedia Rydb., Mem. Torr. Bot. Club 4: 345. 1895; P. monticola 
Mohr, Bull. Torr. Bot. Club 26: 119-120. 1899. 


Plants villous with long jointed hairs, or having only short retrorse ones; 
leaf blades ovate to lanceolate (rarely narrowly so), their margins irregularly 
dentate to sinuate-dentate; corolla usually 15-20 mm. long; anthers yellow, 
or sometimes with a blue or violet tinge. 

TYPE: None cited by Miller. It is supposed to be present in the Sloane 
Herbarium of the British Museum. 

Habitat, distribution and flowering time: Growing in open woods, 
prairies and. disturbed areas in most of the eastern United States, and 
adjacent Canada, extending, generally, into the eastern part of the prairie 
region, with a few collections from the central Rockies; flowering in June 
and July in the northern part of its range, and usually from April to June 


in the southern part. 
(T'o be continued) 


FURTHER NOTES ON THE ILLINOIS FLorA.—Field trips to south- 
ern Illinois in late September and in October, 1957, vielded two 
plants not previously collected in Illinois, a new station for the 
filmy fern in the state and some additional information on the 
height of big bluestem. 

The slender-fruited primrose willow, Jussiaea leptocarpa Nutt., 
ranges from Florida to Texas and Mexico, north to Georgia and 
southeastern Missouri, according to the range given by Fernald in 
Gray's Manual, ed. 8. While botanizing on September 25 in the 


1958] Evers,—Further Notes on Illinois Flora 143 


narrow neck of land known as the Dogtooth Bend in Alexander 
County, I found a few plants of this species! growing on the banks 
of the Mississippi River. On October 24, Dr. M. W. Sanderson, 
an insect taxonomist on the staff of the Illinois Natural History 
Survey, and I visited this site and located numerous individuals 
and a sizable patch of this species. From these plants Dr. San- 
derson collected specimens of a flea beetle of the genus Altica. 
Because of the local abundance of the Jussiaea in this site in the 
Dogtooth Bend, we were curious to learn whether or not it was 
growing farther upstream. Accordingly, we stopped at Fayville, 
a small village opposite Commerce, Missouri, at the mouth of the 
Thebes Gorge of the Mississippi River and approximately 10 miles 
upstream from the Dogtooth Bend location. Here on the banks of 
the river we located some plants. We did not, however, observe 
this species on the riverbank at Thebes, about 5 miles upstream 
from Fayville. The following is a list of specimens of Jussiaea 
leptocarpa Nutt. in the Herbarium of the Illinois Natural History 
Survey (ILLS): 

Riverbank in Dogtooth Bend, south of Miller City, Alexander County, 
Illinois, September 25, 1957, R. A. Evers 55513; October 24, 1957, R. A. 
Evers 55747; riverbank at Fayville, Alexander County, Illinois, October 
24, 1957, R. A. Evers 55781. 

The narrow-leaved sunflower, Helianthus angustifolius L., ranges 
from Florida to Texas and northward to Long Island, New Jersey, 
eastern. Pennsylvania, Kentucky, southern Indiana and south- 
eastern Missouri, according to Gray's Manual. Southern Illinois 
can now be included in the known range. On the northern limits 
of the village of Brookport, Massac County, I located numerous 
individuals of this sunflower and made collections in September 
and October. It was growing in an old field and was associated 
with Bidens sp. and several asters, chiefly Aster pilosus Willd. 
Herbarium sheets were provided with the following data: 

Old field north of Brookport, Massac County, Illinois, September 25, 
1957, R. A. Evers 55625; October 23, 1957, R. A. Evers 55728. 

The filmy fern, Trichomanes boschianum Sturm., which ranges 
from Alabama, north to West Virginia, southern Ohio and southern 
Illinois, is rare in the northern part of its range. Its known range 


1] wish to thank Protessor G. Neville Jones, University of Illinois, for examining this and 
the following species and for verifying my identifications. 


144 Rhodora [ Vor. 60 


in southern Illinois has been restricted to Pope County and based 
upon collections made in only one locality, Jackson Hollow, by 
Mary M. Steagall in 1923, J. W. Swayne in 1950 and R. A. Evers 
in 1951. Plant collectors who botanize in Pope County have been 
seeking this fern elsewhere in the county and also in adjacent 
Johnson County. On October 23, Dr. M. W. Sanderson and I 
were collecting insects and plants in Hayes Creek Canyon, north 
of Eddyville, Pope County, and discovered several small patches 
of T. boschianum on an undercut in a sandstone cliff. Hayes Creek 
Canyon is approximately 5.5 miles east of the Jackson Hollow 
station of this fern. Perhaps a more intensive search in this sec- 
tion of Pope County will reveal other stands of T. boschianum. 
'The herbarium specimen has the following label information: 


Sandstone cliff, Hayes Creek Canyon, north of Eddyville, Pope County, 
Illinois, October 23, 1957, R. A. Evers 55741. 


Thieret and Evers? reported on culms of big bluestem, Andro- 
pogon gerardi Vitman, that measured 3 meters tall. While exam- 
ining a prairie slope on the James M. Nelson property northwest 
of Alton, Madison County, on October 17, Mr. Nelson and I 
observed some culms which we believed to be taller than 3 meters. 
We collected and measured one of the culms and found it to be 3.5 
meters (about 11.5 feet) tall. This culm exceeded by 1.5 meters 
the maximum height for this species as it is described in Hiteheock- 
Chase, Manual of the Grasses of the United States, and by 2 
meters as indicated by Fernald in Gray’s Manual, ed. 8. Ap- 
parently the weather conditions in Illinois in 1957 favored the tall 
and luxuriant growth of big bluestem.—HRonEnT A. EVERS, ILLINOIS 
NATURAL HISTORY SURVEY, URBANA, 


? Ruovora 59 (701): 124. 


Volume 60, number 712, including pages 83-116, was issued 4 June, 1958. 


Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR, 

IVAN MACKENZIE LAMB 


Vol. 60 June, 1958 No. 714 
CONTENTS: 
The Genetic Evaluation of a Taxonomic Character in Dithyrea 
(Cruciferae). ‘Heed C. Rolling... . i. eese aks 145 
A Taxonomic Study of the Genus Physalis in North America North 
of Mexico. U. T. Waterfall (concluded from page 142)....... 152 
X Carex Deamii in Missouri. Julian A. Steyermark............ 174 
An Albino Form of Dipsacus sylvestris. Julian A. Steyermark... 174 
Two New Stations for Carex picta. H.A.Gleason............. 175 
A New Bryophyte Flora. Frank J. Hilferty (Review) ........... 176 


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Vol. 60 June, 1958 No. 714 


THE GENETIC EVALUATION OF A TAXONOMIC 
CHARACTER IN DITHYREA (CRUCIFERAE) 


REED C. ROLLINS 


Developing accuracy in the evaluation of plant characters and 
characteristics for taxonomic purposes is a long-standing prob- 
lem. When differences are found between groups of plants 
which otherwise appear to be related, the immediate question 
arises as to what these differences mean. Specifically, what do 
the differences mean in terms of the genetic make-up of the 
natural group to which such plants belong and how valuable are 
they as taxonomic criteria? In our efforts to interpret speciation 
in relation to a given species or a group of species, we wish to 
rely upon those characters as indicators of relationship (or lack 
of it) that are so deeply seated in the genetic constitution of the 
species that they cannot be easily obliterated or greatly modified 
by the direct effects of any given simply segregating factor or 
combination of segregating factors. In general, the kinds of 
characteristics that offer the greatest possibilities for taxonomic 
reliability are those that are dependent upon a multiplicity of 
genes and gene combinations for their ultimate expression—genes 
that are not in a single linear sequence of interdependency, but 
genes in many series whose interaction in a highly complex way 
results in the final structure or function. It may also be suggested 
that any given characteristic thus dependent upon a complex 
genetical system, which is deeply situated within the genotype, 
becomes protected from radical changes by the build-up of inter- 
dependencies between it and other characteristics, some of which 
may be vital to survival. Thus the species phenotype persists 
over many generations, little altered in basic pattern by the 


146 Rhodora [ Vor. 60 


numerous minor segregations that account for the usual variation 
present. 

At the other end of the scale, characteristics under the control 
of the simplest gene systems are expected to be least reliable as 
the basis for classification. Such characteristics would be easily 
modified or suppressed by repetitious mutations, gene rearrange- 
ments or by ordinary segregation. In consequence, it is probably 
accurate to say that the more simply a character-difference is 
inherited, the less reliable it is as a criterion of speciation. The 
converse of this proposition, that the greater the complexity of 
inheritance of a character-difference the more reliable it is as a 
criterion of speciation, seems equally tenable. In trying to 
evaluate a given genetically controlled characteristic, an im- 
portant attack on the problem is to determine the relative 
complexity of its inheritance. 

One reason that taxonomic characters per se have not been 
frequently subjected to genetic analyses is that this is very time 
consuming and relatively unrewarding. The results of such ex- 
perimental work, though answering the specific question regard- 
ing the nature of the taxonomic character, often do not have 
broader implications. Generalizations can only rarely be made 
because the applicability is or may be restricted to the immediate 
group under investigation. However, it is important for the 
long term to have many more cases worked out than are now 
known. Ultimately these will provide safe guides to proper 
character-evaluations, which is one of the current needs of 
taxonomy. 

In working with the Cruciferae over a period of years, I have 
often encountered situations where "presence" or “absence” of 
trichomes appeared to be of trivial significance (1940, 1952). 
However, without actually testing a given case, it was not 
possible to know whether the absence of an indument in a given 
population, in an otherwise pubescent species, was environ- 
mentally induced or whether it was under genetic control. An 
opportunity to experiment with the presence and absence of a 
dense covering of trichomes on the fruits of Dithyrea Wislizenii 
Engelm. came when a population of this species was found 
having both glabrous- and pubescent-fruited types growing 
together near Sacaton, Arizona. The fruits of individual plants 
of both the glabrous type and the pubescent type were collected 


Rhodora Plate 1233 


PLATE 1233. Flowers and fruits of Dithyrea Wislizenii Engelm. Fig. 1-4, a developmental 
[ I 


series from flower to mature fruit of a pubescent-fruited type. In fig. 1, the petals, 2 sepals 


and the 2 near stamens have been removed to make the ovary visible. The same applies to fig. 


5. Fig. 5-7, a developmental series from flower to mature fruit of a glabrous-fruited type. 


Lard 


1958] Rollins,—Taxonomic Character in Dithyrea 147 


and kept separate for testing purposes. The difference between 
the glabrous siliques and pubescent siliques is very striking, as 
may be seen in Plate 1233. The objective of the following ex- 
periments was to determine the genetic nature of glabrous vs. 
pubescent siliques in this species. 


THe Winp POPULATION 


The species, Dithyrea Wislizenii, extends from western Okla- 
homa and Texas to southern Utah and Nevada, and to Arizona 
and northeastern Mexico. It is common in sandy and loose 
granitie soils and often forms large stands composed of several 
thousands of individuals. Up to the present, a single glabrous- 
fruited Dithyrea, presumably closely related to D. Wislizenii, has 
been recognized as being of some taxonomic worth. Wooton 
and Standley (1913) originally described it at the species level 
as D. Griffithsit and it was later reduced to varietal rank by 
Payson (1918) under D. Wislizenii. In the Sacaton population, 
which provided the material for the following experiments, most 
of the plants possessed pubescent fruits, but there was a goodly 
number of glabrous-fruited individuals. Circumstances did not 
permit a definite count of pubescent vs. glabrous plants in the 
wild population. However, a rough estimate was recorded sug- 
gesting that the pubescent type predominated at least three to 
one. There were no intergrades. The wild plants possessed 
either glabrous fruits or pubescent fruits and none showed a 
gradation from one condition to the other. 

PROGENY TEST OF SEED PARENTS 

Four lots of seeds from the wild population were grown to 
provide plants for crossing purposes. Each seed lot came from 
a single wild plant, which had been open pollinated under natural 


conditions. The plants of culture numbers C-1 and C-4 were 
produced from glabrous-fruited parents, C-2 and C-3 were from 


TABLE I 


WILD PLANTS PROGENIES 
Plant No. Siliques No. Glabrous No. Pubescent 
C-1 glabrous 9 3 
C-2 pubescent 0 15 
7-9 pubescent 0 10 


C-4 glabrous 5 9 


148 Rhodora [Vor. 60 


pubescent-fruited parents. Table I gives the classification of the 
plants of each culture. 

It is of some interest that the progenies of both pubescent 
plants turned out to be uniformly pubescent even though there 
had been no pollen control on the parent plants. 

Pollen mother-cell smears were made to reveal the chromosome 
number of both glabrous and pubescent plants. In each case 
the number n —5 was found. 


CROSSES AND RESULTS 


Three types of crosses were made using various combinations 
from the four cultures originally grown from the wild plants 
listed in Table 1. These were glabrous X glabrous, glabrous X 
pubescent, and pubescent X pubescent. In addition, 14 
pubescent plants were placed together in an isolated greenhouse 
where interpollination was permitted to be effected by the 
insects normally present. In each of the three types of controlled 
crosses, bagging with muslin, emasculation and hand pollination 
were practiced. Controls to check the procedures were carried 
along with the experiments. These showed that pollen control 
was effective. All crosses were carried out reciprocally. Essen- 
tially the same results were achieved regardless of the direction 
in which the pollen was carried except for the reciprocal of 


TABLE 2. GLABROUS X GLABROUS 


CROSS PROGENY 3:1 RATIO CHI-SQUARE 
No. No. No, 
plants glabrous pubescent 
Cl-1 x Cl1-3 17 12 5 
reciprocal 7 5 2 
24 17 7 18:6 .16 
Cl-4 x C4-9 20 15 5 
reciprocal 6 4 2 
26 19 7 19.5:6.5 .05 
Cl-7 x C4-7 12 9 3 
reciprocal 8 7 1 
20 16 4 15:5 .266 
Total 70 52 18 287 


P = .98-.95 


1958] Rollins,—Taxonomic Character in Dithyrea 149 


pubescent C1-59 X glabrous Cl-7%. In this case, the 
reciprocal did not produce any filled seeds. The significance of 
this failure was not determined. 

In addition to the results shown in Tables 2, 3 and 4, fourteen 
progenies of pubescent plants open pollinated from pubescent 
plants were grown. These amounted to 159 plants, all of which 
possessed pubescent siliques. 


TABLE 3. GLABROUS X PUBESCENT 


CROSS PROGENY 1:1 RATIO CHI-SQUARE 
No. No. No. 
plants glabrous pubescent 


C1-7 x C1-5 0 0 0 
reciprocal 10 3 7 
10 3 7 5:5 1.60 
C1-6 x C4-8 12 3 9 
reciprocal 11 4 7 
23 7 16 11.5:11.5 3.52 
C4-13 x C4-6 6 4 2 
reciprocal 14 5 9 
20 9 11 10:10 .20 
C4-10 x C4-14 17 6 11 
reciprocal 6 3 3 
23 9 14 11.5:11.5 1.08 
Total 76 28 48 6.40 
P = .2-.1 
Pooled Chi-square (1 df) 5.26 
P = .05-.02 
Heterogeneity Chi-square (3 df) 1.14 
P = 3-7 


The results are easily explainable if it is assumed that a single 
gene pair is operative in producing the glabrous or pubescent 
condition of the siliques. From the data, it is obvious that the 
pubescent plants are homozygous and recessive. Thus the geno- 
type of the pubescent plants may be designated gg. When such 
a plant is crossed with a glabrous heterozygous individual (Gq), 
the resulting progeny should show a 1:1 ratio of glabrous to 
pubescent plants. In table 3, results from four different crosses 
between glabrous and pubescent plants are given and the Chi- 
square test for goodness of fit to a 1:1 ratio is provided. The 


150 Rhodora [ Vou. 60 


TABLE 4. PUBESCENT X PUBESCENT 


CROSS PROGENY 
No. glabrous No. pubescent 

C2-7 x C2-8 0 11 
reciprocal 0 1 
0 12 
C3-2 x C3-9 0 10 
reciprocal 0 8 
0 18 
C3-6 x C3-7 0 13 
reciprocal 0 28 
0 41 
C4-11 x C4-12 0 17 
reciprocal 0 30 
0 47 
Total 0 118 


numbers of plants in the various progenies are small and the 
possibility of results different from those shown should perhaps 
not be ruled out completely. However, the evidence strongly 
favors a 1:1 ratio and the assumption of a heterozygous (Gg) 
plant as the glabrous parent in each cross seems justified. 

If heterozygotes are crossed, a 3:1 ratio of glabrous to pubescent 
is to be expected. Table 2 gives the data on three glabrous X 
glabrous crosses and the results show convincingly that a 3:1 
ratio of glabrous to pubescent was obtained. It seems perfectly 
safe to assume that each of the six parents was of the constitution 
Gg with respect to the genes in control of the glabrous vs. pubes- 
cent condition. Evidently no homozygous dominant plants were 
used in the experiments. Such plants could not be distinguished 
from the heterozygotes phenotypically. 


DISCUSSION 

The mechanism of genetic control of glabrous vs. pubescent 
siliques in Dithyrea Wislizenit is obviously a relatively simple 
one. For this reason, it is safe to reject the phenotypic charac- 
teristic of glabrous siliques as having no significance for taxo- 
nomic purposes. Plants with this characteristic are expected to 
occur without respect to phylogenetic relationship in the popula- 
tions of the species. In fact, this is exactly what one finds. 


1958] Rollins,—— l'axonomie Character in Dithyrea 151 


Glabrous-fruited plants are found in D. Wislizenii proper and in 
D. Wislzenii var. Palmeri. Furthermore, glabrous-fruited 
plants are found more or less throughout the geographical range 
of the species. On the basis of collections in the Gray Her- 
barium, the pubescent type appears to be more common than 
the glabrous. Collectors evidently distinguish between the 
glabrous and pubescent plants in the field because there are but 
three mixed collections among 84 different ones available in the 
herbarium. Six of the collections have glabrous fruits and 75 
have pubescent fruits. 

In my own field experience, I have examined four different 
populations of D. Wislizenii, one in Texas, two in Arizona and 
one in New Mexico. In three of these populations, I was unable 
to locate any glabrous-fruited plants. The fourth population 
vielded the material reported on above. This evidence added to 
that from herbarium material makes it quite clear that pubescent- 
fruited plants predominate in the species. If pubescent siliques 
represent the homogygous recessive condition, as indicated by 
the analysis of the one population, a more prevalent occurrence 
of the glabrous type would be expected throughout the species 
as a whole unless there are positive factors operating to select 
against it. However, we have no way of knowing about this at 
the present time. One observation may be pertinent to any 
ultimate explanation for the prevalence of the pubescent-fruited 
type. It is that the plants of D. Wislizenti are self incompatible. 
The chances of the accidental establishment of predominantly 
glabrous-fruited populations through isolation are considerably 
reduced as compared with a self compatible species. 


CONCLUSIONS 


The glabrous-fruited condition in Dithyrea Wislizenti, which 
provided the chief basis for describing D. Griffithsii Wooton and 
Standley as a separate species, is a simply inherited characteristic 
under single gene control. Glabrous-fruited heterozygotes when 
crossed produce a simple mendelian 3:1 ratio of glabrous- to 
pubescent-fruited plants. Glabrous-fruited heterozygotes 
crossed with pubescent-fruited plants produce approximately a 
1:1 ratio of glabrous to pubescent plants. Pubescent-fruited 
plants crossed with each other produce only pubescent-fruited 
offspring. The pubescent plants studied all proved to be 


152 Rhodora [ Vor. 60 


homozygous and recessive for the pubescent fruit character. 
The presence or absence of pubescence on the siliques of D. 
Wislizenii is of no taxonomic significance. 
—GRAY HERBARIUM OF HARVARD UNIVERSITY. 

LITERATURE Crrep 
Payson, E. B. 1918. Notes on Certain Cruciferae. Ann. Mo. Bot. 

Gard. 5: 148. 

Ronums, Reep C. 1940. A Monographie Study of Arabis in Western 


North America. RHODORA 43: 299. 
—,. 1952. Taxonomy Today and Tomorrow. RHODORA 


54: 13. 
Wooron, E. O. and P. C. SraNpLEvy. 1913. Contrib. U. S. Nat. Herb. 
16: 124. 


A TAXONOMIC STUDY OF THE GENUS PHYSALIS 
IN NORTH AMERICA NORTH OF MEXICO 


U. T. WATERFALL 
(Concluded from p. 142 


Among the 481 sheets of 450 collections studied, material has been 
seen from Manitoba, Ontario (Macoun 54525, Ny) and Quebec (Marie- 
Victorin et al 46421, au) in Canada, and in the United States from: Ala- 
bama, Arkansas, Colorado (Boulder and El Paso Cos.), Connecticut, 
Delaware, Florida (Lafayette Co.), Georgia, Illinois, Indiana, Iowa, Kan- 
sas, Kentucky, Louisiana (Short, NY), Maine (Cumberland Co.), Massachu- 
setts, Miehigan, Minnesota, Mississippi (University Campus), Missouri, 
Nebraska, New Hampshire (Coós and Merrimack Cos.), New Jersey 
(Middlesex Co.), New York (Staten Island), North Carolina, North 
Dakota, Ohio (Lorain Co.), Oklahoma, Pennsylvania (Chester Co.), 
Rhode Island, South Carolina (Aiken Co.), South Dakota, Tennessee, 
Texas, Utah (Washington Co.) Virginia, Washington D. C., West Vir- 
ginia (Monroe Co.) and Wisconsin. 

9b. P. virginiana Miller, var. subglabrata (Mackenzie and Bush) 
Waterfall, comb. et stat. nov., based on P. subglabrata Mackenzie and 
Bush, Trans. Acad. Sci. St. Louis 12: 86-87. 1902. 

Plants nearly glabrous, or with a few short antrorse hairs; leaf blades mostly 
ovate to ovate-lanceolate, their margins usually entire, sometimes slightly 
sinuate-dentate; anthers tinged or margined with blue or violet; fruiting calyces 
mostly 25-35 mm. long and 20-30 mm. wide. 

TYPE: K. K. Mackenzie collected at Sheffield, Jackson Co., Missouri, 
June 14, 1896 (xv). 

This variety grows in woods, grassland, roadsides, fields, and other dis- 
turbed sites, primarily in the northeastern United States, but with scat- 
tered eolleetions elsewhere; it flowers mostly from June to September. 


1958] Waterfall—Genus Physalis in N. America 153 


Four hundred thirteen sheets of 267 collections have been studied. This 
taxon oceurs in Ontario, Canada, and in Arkansas, Colorado (Gunnison 
and Routt Cos.), Connecticut, Delaware, Georgia (Wayne Co.), Idaho 
(Ada, Canyon and Payett Cos.), Illinois, Indiana, Iowa, Kansas (Douglas, 
Ellis and Shawnee Cos.), Kentucky, Louisiana (East Feliciana and Grant 
Parishes), Maryland, Massachusetts, Michigan, Mississippi (Coahoma 
Co.), Missouri, Nebraska (Kearney and Nemaha Cos.), New Jersey, New 
Mexico (Lincoln Co.), New York, North Carolina, Ohio, Oklahoma 
(Rogers and Tulsa Cos.), Oregon (Polk Co.), Pennsylvania, Rhode Island, 
South Carolina (Pickins Co.), Tennessee, Texas (Delta, Nueces and Potter 
Cos.), Utah (San Juan and Sanpete Cos.), Virginia, Washington D. C., 
West Virginia, Wisconsin. 

Sometimes forms are found with larger fruiting calyces which are 4-5 
em. long and 3-4 em. broad. These may be called Physalis virginiana 
f. macrophysa (Rydberg) Waterfall, comb. et stat. nov. based on P. 
macrophysa Rydberg, Bull. Torr. Bot. Club 22: 308. 1895. 

TYPE: Since Rydberg cited several number without choosing a type, the 
author selects as LEcrorvPE A. A. Heller 1756 (NY); isolectotypes: (NY, 
UC). 
Although the latter name antedates subglabrata, the author has chosen 
to transfer the name subglabrata to varietal status, since it is associated 
with a large, wide-spread population, primarily of the northeastern United 
States. According to Article 70 of the Rules no name has priority outside 
its own rank. Article 71, Recommendation 71A suggests that in changing 
rank it is preferable to retain the original epithet unless it must be rejected 
under the rules. This particular choice of names seems to be in accord- 
ance with both articles, and keeps available the names now in usage, in- 
stead of either changing their application, or supplanting them with new 
names which would be permissible under the rules. 

Selected specimens: ARKANSAS Marion Co.: Demaree 20645 (NY); 
Inuivois: Peoria Co.: McDonald Aug. 1903 (Ny); Champaign Co.: Gleason 
Oct. 7, 1907 (puxe); Inprana: Lawrence Co.: Kriebel 1348 (pvKE); Iowa: 
Story Co.: Hayden 424 (au); Missouri: Jackson Co.: Bush 12488A (NY) 
and 12483 (DUKE); NeBraska: Howard Co.: Bates 4910 (au); New Jersey: 
Somerset Co.: Lighthipe Aug. 1, 1916 (Tex); Texas: Comal Co.: Lind- 
heimer May 1847 (au); Kerr Co.: Heller 1756 (Ny, uc); Tarrant Co.: Ruth 
746 (xy, Pu); Travis Co.: Tharp May 6, 1931 (rex). 

Oc. P. virginiana Miller, var. texana (Rydberg) Waterfall, comb. et 
stat. nov., based on P. texana Rydberg, Mem. Torr. Bot. Club 4: 339—340. 
1896. 

Plant usually several-branched from the base; herbage glabrous or nearly 
so; principal leaves ovate and usually entire; plant of the Gulf coast of Texas, 
apparently intergrading inland with contiguous varieties. 

TYPE: The type is A. A. Heller 1507 Ny; isotypes: ARIZ, GH, PH, UC. 

This is primarily a taxon of the coastal area, but it extends inward 
through chaparral and other habitats nearly to central Texas, becoming 
more atypical as it does so. It usually flowers from March to June, but 


154 Rhodora [ Vor. 60 


flowering specimens have been seen that were collected in December and 
in August. 

Selected from 35 sheets of 17 collections: Texas: Bexar Co.: Metz 73 
(micu, NY); Cameron Co.: Parks 17943 (aH); Gonzales Co.: Tharp 61-556 
(rex); LaSalle Co.: Tharp and Tyson 52-488 (okta, TEX); Nueces Co.: 
Tharp, Johnston and Webster 48-58 (TEX, ARK, OKLA); San Saba Co.: 
Palmer 11841 (vex); Travis Co.: Tharp and Scarbrough 51-399 (coro, 
OKLA, TEX, UARK); Victoria Co.: Tharp 2516 (rex); Washington Co.: 
Tharp July 9, 1929 (oxra, TEX); Wilson Co.: Palmer 947 (Gu, NY). 

Od. P. virginiana Miller, var. sonorae (Torrey) Waterfall, comb. nov., 
based on P. pumila Nutt., var. sonorae Torr., Botany of the Mexican 
Boundary 153. 1859; P. longifolia Nutt., Trans. Am. Phil. Soc. (n.s.) 5: 
193-194. 1836; P. lanceolata Michx., var. laevigata Gray, Proc. Am. Acad. 
Arts and Sciences 10: 68. 1875; P. lanceolata Michx., var. longifolia 
(Nutt.) Trelease, Rep. Ark. Geol. Surv. 4: 207. 1891; P. rigida Pollard 
and Ball, Proc. Biol. Soc. Wash. 13: 134-135. 1900. 

Since, according to Article 70, “When the rank . .. of an infragenerie 
taxon is changed, the correct name or epithet is the earliest legitimate one 
available in the new rank," the well-known name longifolia must be re- 
placed in the varietal status with the relatively unknown sonorae. 

Plants usually single stemmed, often branching above; leaf blades usually 
lanceolate to lanceolate-lirear, but rarely ovate, their margins entire to irregu- 
larly toothed; herbage sparsely covered with short antrorse hairs, which are 
more abundant on the younger parts, sometimes nearly glabrous; calyx often 
with ten lines of short antrorse hairs; anthers yellow. Sometimes this variety 
is difficult to separate from var. subglabrata. In such cases the bluish, or 
violet, anthers of the latter is considered a distinguishing characteristie since 
it occurs in a large population of the northeastern United States where the 
vellow-anthered var. sonorae is not found. 

TYPE: Geo. Thurber 418, Fronteras, Sonora, Mexico, June 1851 in the 
Herbarium of the New York Botanical Garden. Two isotypes are in the 
Gray Herbarium. 

Habitat, distribution and flowering time: This variety grows in prairies, 
plains, foothills, canyons, open woods, sandy areas and in various disturbed 
habitats. Its primary distribution is in the prairie region of central 
United States, but it extends into and west of the Rockies. It flowers in 
June, July and August in the northern part of its range, and from May 
through September in Texas and Arizona. 

Four hundred sixty seven sheets of 385 collections have been studied. 
They were collected in Arizona, Arkansas, California (Trinity Co.), Color- 
ado, Georgia (Whitfield Co.), Idaho, Illinois, Indiana, Iowa, Kansas, 
Nebraska, Nevada (Churchill and Storey Cos.), New Mexico, North 
Carolina, Oklahoma, Oregon (Malheur Co.), Pennsylvania (Small, 1889), 
South Dakota, Tennessee (Davidson Co.), Texas, Utah, Virginia (King 
George Co.), West Virginia (Mineral Co.), Wisconsin, and Wyoming 
(Park and Weston Cos.). 

It is probable that the records from the far west represent introductions. 

9e. P. virginiana Miller, var. hispida Waterfall, var. nov., foliis ovato- 


1958] Waterfall,—Genus Physalis in N. America 155 


lanceolatis vel lanceolatis, vel spathulato-lanceolatis, crassis, plus minusve 
hispidis. 

This perennial rhizomatous variety has thick leaf blades varying from 
ovate-lanceolate to linear-lanceolate in shape. The herbage is sub- 
glabrous, but it has a varying amount of stiff, more or less divergent 
trichomes about 1 mm. long, at least on the flower buds or the margins of 
the leaves. It is found in sandy areas of the prairie and plains region. It 
has been passing as P. lanceolata, but that name was given to seemingly 
aberrant plants of the eastern United States which may very well be inter- 
grades between P. heterophylla and P. virginiana, 

The type is Waterfall 7308, sand dunes, 1 mile east of Mangum, Greer 
Co., Oklahoma, June 28, 1947, in the Herbarium of Oklahoma State 
University; isotypes (TEX, OKL). 

It usually grows in sandy situations, but is found also on dry hilltops, 
edges of fields and other disturbed areas, primarily in Oklahoma, Kansas, 
Nebraska and eastern Colorado; flowering in May, June and July. 

Selected from 212 sheets of 147 collectiors: Cotorapo: Baca Co.: Rogers 
6436 (cono); Boulder Co.: Ramaley 11624 (coLo); Denver Co.: Eastwood 23 
(COLO, GH, vc); El Paso Co.: Williamson July 10, 1901 (pH); Fremont Co.: 
Brandegee 392 (NY, PH, vc); Jefferson Co.: Greene 323 (an); Larimer Co.: 
Smith July 15, 1944 (vc); Las Animas Co.: Rogers 6003 (coro); Lincoln Co.: 
Ownbey 1318 (cono, GH, NY, vc); Phillips Co.: Weber 5040 (coro); Weld Co.: 
Ramaley 15138 (ARIZ, COLO, OKL, TEX, UC); Yuma Co.: Harrington 5036 (COLO); 
InprANa: Lake Co.: Bebb 499 (oKL); Tippecanoe Co.: Ek June 10, 1942 (an, 
NY, TEX, UC); Kansas: Barton Co.: Rydberg and Imler 1330 (KANU, NY); 
Cheyenne Co.: McGregor 9433 (Kanu); Clark Co.: Rydberg and Imler 768 
(KANU); Clay Co.: Kellerman July 2, 1888 (GH); Comanche Co.: Rydberg and 
Imler 1109 (KANU, NY); Finney Co.: Rydberg and [mler 996 (KANU, NY); Grove 
Co.: Hitchcock 572 (Gu); Hamilton Co.: Wilson and Miller (kanu); Meade Co.: 
Horr and McGregor 3841 (Kanu); Reno Co.: Rydberg and Imler 563 (KANU, 
nY); Riley Co.: Norton 368 (an, NY); Rooks Co.: Horr 5005 (Kanu); Sedgwick 
Co.: Coll. unknown Aug. 20, 1933 (kanu); Wyandotte Co.: Mackenzie 1159 
(NY); MiNNESOTA: Isanti Co.: Rosendahl and Butters 5051 (Gu); Missouri: 
Jackson Co.: Bush 4970 (Gu, Ny, OKL); Johnson Co.: Stevens 4166 (ny); Ne- 
braska: Banner Co.: Rydberg 473 (NY); Cherry Co.: Tolstead 550 (Gu); Custer 
Co.: Bates June 15, 1901 (Gu); Hapeman June 7, 1928 (puKE); Lincoln Co.: 
Porter 2059 (Gu, okL); New Mexico: Santa Fe Co.: Tracy and Evans 110 
(NY); OKLAHOMA: Beckham Co.: Pennell 10556 (Ny, pH); Beaver Co.: Goodman 
5332 (OKL, TEX); Blaine Co.: Waterfall 7070 (OKL, OKLA, TEX); Cleveland Co.: 
Little 396 (oKL); Custer Co.: Mericle 318, 328, 703, 1866 (okr); Ellis Co.: 
Waterfall 11891 (oKraA); Garvin Co.: Andrews 97 (OKL); Grady Co.: McFarland 
15 (OKL); Greer Co.: Bull 219 (okr); Harper Co.: Stevens 3322 (GH, NY, OKL); 
Jackson Co.: Stevens 1176 (GH, OKL); Kay Co.: Stevens 1919 (Gu); Kingfisher 
Co.: Byers 211 (oKLA); Logan Co.: Goodman 2126 (GH, OKL); Oklahoma Co.: 
Waterfall 2350 (Gu, NY); Payne Co.: Abernathy 32 (oxra); Pushmataha Co.: 
Waterfall 11397 (OKLA); Roger Mills Co.: Smith 607 (oKLA); Texas Co.: Water- 
fall 7961 (OKL, OKLA); Woods Co.: Waterfall 7857 (OKL, OKLA); Woodward 
Co.: Nelson and Goodman 5301 (oKL); South Dakota: Meyer Co.: Wallace 
(NY); Texas: Collingsworth Co.: Cory 16151 (au); Hall Co.: Reverchon 4511 
(Gu); Hemphill Co.: Cory 16236 (Gu); Wichita Co.: Tharp 535 (wy, TEX); Uran: 
Sanpete Co.: Ward 676 (Gu); Wyomina: Albany Co.: Nelson 7358 (GH, NY); 


jt 


156 Rhodora | Vor. 60 
Converse Co.: Nelson 8366 (an); Platte Co.: Porter 4894 (COLO, GH, OKL, PH, 
TEX, UC). 

Of. P. virginiana Miller, var. polyphylla (Greene) Waterfall, comb. et 
stat. nov., based on Physalis polyphylla Greene, Pittonia 4: 150-151. 
1900. 

Plants simple or branched near the base, nearly glabrous, the few hairs 
short and antrorse; longer leaves mostly 3-5 cm. long, lanceolate or 
linear-lanceolate. 

type: C. F. Baker 576 Piedra, southern Colorado, July 12, 1899 (iso- 
types: GH, NY, UC). 

The only other collection seen is Waterfall 11115, collected on a shale 
hillside, opening in pine forest 12 miles west of Chama, Rio Arribo Co., 
New Mexico, Aug. 25, 1952. (OKLA). 

Og. P. virginiana Miller, var. campaniforma Waterfall, var. nov., 
caulibus parvis, eurtis retrorso-pilosis; foliis ovatis; corollis maculatis; 
‘alycis eampaniformis, ad basin 4-5 mm. latis, ad apices 1.5-2 em. latis. 

This variety is characterized by the combination of ovate leaves, short 
retrorse hairs and campanulate calyx, 4-5 mm. wide at its base, and 1.5-2 
em. wide at the tips of its divergent lobes. 

TYPE: P. C. Standley 4556. Mouth of Indian Creek, altitude 8000 ft. ; 
in Pecos National Forest, New Mexico, July 25, 1908. Two sheets are 
in the Herbarium of the New York Botanical Garden. At present this 
distinctive variety is known only from the type collection. 

10. Physalis hederaefolia Gray, Proc. Amer. Acad. Arts and Sciences 
10: 65. 1875. 

Plants erect or spreading from a perennial base, simple or many-stemmed; 
herbage with a mixture of long jointed hairs and short trichomes, or with short 
hairs only, which may be viscid or not, glandular or not, antrorse or spreading; 
leaf blades subreniform to ovate, or rarely ovate-lanceolate; corollas 10-15 
mm. long, yellow or yellowish green, usually darker on the base of the limb, 
but sometimes obscurely so; limb of the corolla often reflexed when fully open; 
anthers usually vellow, 1.5-4 mm. long; flowering calyx about one-half as long 
as the corolla, on peduncles usually 3-8 mm. long; fruiting calyx 2-3 em. long 
and 1.5-2.5 em. wide on peduncles 1-2 cm. long. 

10a. P. hederaefolia Gray, var. hederaefolia P. hederaefolia Gray, var. 
puberula Gray, loc. cit. supra; P. Palmeri Gray, Synoptic Flora 2 (1): 235. 
1588. 

Herbage vestite with one or some combination of long jointed hairs, short di- 
varicate or retrorse hairs, or glandular hairs; leaf blades subreniform to ovate; 
calyces at anthesis usually 3-4 mm. wide; anthers mostly 3-4 mm. long, yellow 

TYPE: Charles Wright 528, in part, Turkey Creek, western Texas to El 
Paso, Oct. 1849 (au). It has a few long hairs, a few short hairs and a few 
sessile, or subsessile spherical glands; the type of var. puberula has many 
short hairs on the stems, and short hairs and a few spherical sessile glands 
on the leaves. 

Variety hederaefolia grows in desert plains, desert scrub, canyons, 
mountains and valleys, primarily in southwestern Texas, New Mexico and 
adjacent Arizona; it flowers in May, June and July, sometimes in August, 
September and October. 


1958] Waterfall, —Genus Physalis in N. America 157 


Selected from 243 sheets of 180 specimens: ARIZONA: Cochise Co.: Harrison 
8259 (ariz); Coconino Co.: Hanson 133A (coro, TEX); Gila Co.: Gould and 
Hudson 3742 (ariz, GH, Uc); Pima Co.: Gould 3952 (ariz); Pima Co.: Toumey 
404 (artz); Yavapai Co.: Wolf 2311 (au); CALIFORNIA: San Bernardino Co.: 
Wolf 10772 (vc); San Diego Co.: Abrams 3703 (an); Nevapa: Clark Co.: 
Clokey 8107 (au, NY); Lincoln Co.: Ripley and Barneby 6405 (NY); NEw 
Mexico: Bernalillo Co.: Koelz June 28, 1926 (urcn); Dona Ana Co.: Wooton 
136 (NY); Grant Co.: Greene Sept. 31, 1880 (Ny); Lincoln Co.: Wooton 634 (NY); 
San Miguel Co.: Rose and Fitch 17606 (Ny); Texas: Cory 16628 (au); Bandera 
Co.: Palmer 12253 (tex); Brewster Co.: Mueller 8154 (GH, MICH, NY, TEX, UC); 
Warnock 341 (GH, NY, TEX); Burnet Co.: Rogers, Albers and Barksdale 6864 
(OKLA, TEX); Cameron Co.: Chandler 7064 (GH, NY, vc); Culberson Co.: Water- 
fall 4059 (Gu, NY); Duval Co.: Croft 11 (micu, Ny); El Paso Co.: Warnock 4100 
(rex); Gonzales Co.: Smith and LeSueur 42-42 (an, TEX); Grimes Co.: Tharp 
April 11, 1936 (rex); Hidalgo Co.: LeSueur 442 (rex); Hudspeth Co.: Waterfall 
4875 (an, NY); Jeff Davis Co.: Palmer 31935 (TEx); Kenedy Co.: Cory 28408 
(au); Leon Co.: Cory 21810 (au); Mason Co.: Whitehouse Sept. 1, 1929 (TEX); 
McLennan Co.: Smith 620 (TEx); Maverick Co.: Pringle 8324 (GH, NY, UC); 
Pecos Co.: Tharp 256 (okr, vc); Presidio Co.: Hinckley 2753 (au); Real Co.: 
Cory 42778 (au); Reeves Co.: Tracy and Earle 126 (an, NY, TEX); Smith Co.: 
Cory 25881 (au); Taylor Co.: Cory 7393 (micu, vc); Terrell Co.: Webster 190 
(TEX); Tom Green Co.: Reverchon 3922 (Gn); Travis Co.: Tharp Aug. 18, 1941 
(au, TEX); Upton Co.: Cory 53482 (cu); Webb Co.: Mackenzie 86 (Ny); UTAH: 
Kane Co.: Boyle 208 (uc); Millard Co.: Garrett 2969 (Ny). 

10b. P. hederaefolia Gray, var. comata (Rydberg) Waterfall, 
Ruopora 52: 171. 1950; P. comata Rydb., Bull. Torr. Bot. Club 22: 306. 
1895; including P. rotundata Rydb., Mem. Torr. Bot. Club 4: 352. 1896. 


Herbage with long jointed hairs more or less abundantly mixed with shorter 
hairs, which may, or may not, be viscid or glandular; flowering calyx 8-11 mm. 
wide; leaf blades ovate to rotund, toothed to nearly entire. 

TYPE: P. A. Rydberg 269, under the cliffs, south side of Scott's Bluff, 
Nebraska, July 20, 1891, in the Herbarium of the New York Botanical 
Garden. 

Variety comata is found on plains, mountains slopes, dry hills, gravel 
banks and sandhills, principally in western Nebraska, western Kansas and 
eastern Colorado; it usually flowers in June, July, August and September. 


Selected from 80 sheets of 63 collections: CoLorapo: Bent Co.: Osterhout 
4118 (Ny); Boulder Co.: Ewan 12258 (uc); Cheyenne Co.: Ownbey 1357 (coro, 
an, Ny); Denver Co.: Eastwood Sept. 10, 1910 (au, vc); El Paso Co.: Ehlers 
7764 (ARIZ, GH); Fremont Co.: Ewan 14248 (coro); Larimer Co.: Nelson Aug. 
31, 1900 (xv); Las Animas Co.: Rogers 4843 (coro); Weld Co.: Osterhout 2309 
(xy); Kansas: Barber Co.: Rydberg and Imler July 5, 1929 (ny); Cheyenne 
Co.: McGregor 9425 (Kanu); Ellis Co.: Rydberg and Imler 1228 (KANU, NY); 
Finney Co.: Wilson and Miller July 22, 1912 (KANv); Kiowa Co.: Hitchcock 774 
(aH, NY); Osborne Co.: Shear 221 (an, Ny); Riley Co.: Gates 14537 (MICH); 
Rooks Co.: Bates 4563 (au); NEBRASKA: Adams Co.: Bates 4607 (au); Rydberg 
Aug 8, 1891 (xv); Buffalo Co.: Bates 4903 (au); Custer Co.: Bates 2403 (an); 
Garfield Co.: Bates 4634 (ny); Lincoln Co.: Rydberg Sept. 1895 (uc); Webster 
Co.: Bates July 3, 1907 (xv); New Mexico: Lincoln Co.: Skehan July 7, 1898 
(GH, NY, UC); OkLAHOMA: Cimarron Co.: Waterfall 7902, 9122, 9240 (OKL, 
OKLA); Texas Co.: Waterfall 7867 (OKL, OKLA, TEX); Woods Co.: Ward 54 
(xy); Soura Dakota: Lawrence Co.: Bennett 3259 (UARK). 


158 Rhodora [ Vor. 60 


10c. P. hederaefolia Gray, var. cordifolia (Gray) Waterfall, eomb. 
nov., based on P. Fendleri Gray, var. cordifolia Gray, Synop. Flora N. 
Amer. 2(1): 395. 1878. P. Fendleri Gray, Proc. Amer. Acad. Arts and 
Sciences 10: 66. 1875. 

Indument of short reflexed hairs with many to few short, somewhat 
flattened branched hairs, which are sometimes present onlv on the calyx; 
leaf blades ovate to ovate-lanceolate (2) 3-6 em. long and 1-3 em. wide, 
from (1.2) 1.4-2.5 times longer than wide. 

TYPE: The type of P. Fendleri is Fendler 683, New Mexico (au). That 
of var. cordifolia is Palmer 363, St. George, southern Utah (an). 

This taxon grows in mountains, canyons, mesas, plains, in juniper-pinon 
pine areas, and in disturbed habitats, principally in southwestern Texas; 
it flowers mostly in July, August and September. 


Selected from 196 sheets of 146 collections: Arizona: Apache Co.: Goodman 
and Payson 3167 (an, NY); Cochise Co.: Blumer 2104 (Ariz, GH, NY); Coconimo 
Co.: Thornber 2088 (ariz); Gila Co.: Parker, McClintock and Robbins 6125 
(ariz); Maricopa Co.: Rusby 775 (mich, Ny); Mohave Co.: Kearney and 
Peebles 12761 (ariz); Navajo Co.: Jones 1109 (ariz); Pima Co.: Parker, 
McClintock and Haskell 5885 (ariz, vc); Santa Cruz Co.: Peebles and Loomis 
7019 (artz); Yavapai Co.: Kearney and Peebles 9719 (ARIZ); CALIFORNIA: 
San Bernardino Co.: Wolf 10722 (ny, uc); San Diego Co.: Palmer 1875 (au); 
Cotorapo: Archuleta Co.: Weber and Livingston 6254 (coto); El Paso Co.: 
McCosh and Greene 1877 (Ny); Fremont Co.: Waterfall 11503 (OKLA, TEX); 
Huerfano Co.: Ramaley 16236 (coto); La Plata Co.: Jones 503 (micu); Las 
Animas Co.: Rogers 5416 (COLO); Mesa Co.: Rollins 1915 (Gu, NY); Montezuma 
Co.: Baker, Earle and Tracy 823 (ny); Montrose Co.: Payson 3927 (aH); 
Otero Co.: Paull 8? (cono); Pueblo Co.: Pammel Aug. 24, 1913 (an, TEX); 
Nevapba: Clark Co.: Train 2003 (ariz); Clokey 8204 (ARIZ, DUKE, NY, OKL, 
OKLA, TEX); New Mexico: Colfax Co.: Standley 14012 (ny); Dona Ana Co.: 
Wooton and Standley 3157 (ariz, Ny); Grant Co.: Blumer 49 (au, NY); Luna 
Co.: Shreve 8343 (ariz); Otero Co.: Schulz 297 (Gu); Fan Miguel Co.: Standley 
4945 (GH, NY); Santa Fe Co.: Robbins 8244 (coro); Sierra Co.: Metcalfe 945 
(GH, NY); Taos Co.: Wooton 2693 (Ny); Torrance Co.: Parker and McClintock 
6526 (arız); Valencia Co.: Vogt 27 (ariz); Oklahoma: Cimarron Co.: Waterfall 
7915 (OKL, OKLA); Texas: Brewster Co.: Marsh 261 (au); El Paso Co.: Lee, 
Berkman and Tharp 46192 (Tex); Hudspeth Co.: Waterfall 6694 (au); Jeff 
Davis Co.: Hinckley 574 (xy); Uran: Piute Co.: Tidestrom 2942 (micah); San 
Juan Co.: Rydberg and Garrett 9390 (NY); Washington Co.: Gould 2028 (ARIZ, 
COLO, GH, NY). 

11. Physalis caudella Standley, Field Mus. Publ. Bot. 17: 273. 1937. 

Plants simple or branched, apparently from a deep rhizome which is not 
collected; indument usually villous, of long jointed hairs (1) 2-3 mm. long, 
dense or sparse, or of long and short hairs intermixed in varying proportions; 
leaf blades 4-7 em. long and 1.5-4 em. wide, usually lanceolate, rarely ovate- 
lanceolate or linear-lanceolate, on petioles 0.5-2 em. long (this amount of 
variation in length of petioles may be found in the same plant, with the longer 
petioles below and the shorter ones above); margins of the leaf blades entire 
to irregularly undulate to saliently few-toothed; corollas 14-18 mm. long, 
yellow, with prominent deep reddish-blue or purplish spots on the limb; 
anthers blue or blue-green, about 3 mm. long, on slender filaments much 


1958| Waterfall, —Genus Physalis in N. America 159 


narrower than the anthers; calyx 7-10 mm. long, its lobes 3-8 mm. long; 
flowering peduncles usually about 5 mm. long, sometimes as much as 8 mm. 
long; fruiting calyx (2.5) 3-5 em. long and (2) 2.5-3 cm. wide, with calyx 
lobes (6) 10-15 (17) mm. long. 

TYPE: Howard Scott Gentry 2710, on oak-pine slope, 2,160 meters eleva- 
tion, Cajurichi, Rio Mayo, Chihuahua, Mexico, Sept. 13, 1936 (r). An 
isotype is at uc. The isotype has lobes of the flowering calyx 3-8 mm. 
long, and a calyx cup only about 2 mm. long; the fruiting calyx also has 
lobes at the extreme limit of length, being 14-17 mm. long. One of the 
Arizona specimens approaches the type, having a flowering calyx with 
lobes 7 mm. long and a calyx tube 3 mm. long. However most of them 
have ealyx lobes somewhat shorter than the calyx tube; the lobes of the 
fruiting calyx in the Arizona material are usually 10-15 mm. long, rarely 
as short as 6 mm. 

Habitat, range and flowering time: Growing in canyons, pine woods and 
oak woods in the mountains of southern Arizona (with one collection from 
southwestern New Mexico) and adjacent Sonora and Chihuahua; flowering 
in June, July and August. 

Selected from 31 sheets of 19 collections: Arizona: Cochise Co.: Benson 
10448 (Antz, NY, UC); Gooding 843 (arız, GH, NY); Pima Co.: Kearney and 
Peebles 10504 (ariz, uc); Santa Cruz Co.: Parker 7683 (ARIZ, COLO, NY, 
uc); New Mexico: Socorro Co.: Wooton Aug. 6, 1900 (NY). 

12. Physalis crassifolia Bentham, Botany of the Voyage of the Sulphur 
40. 1844. 

Stems usually several from a ligneous base, each stem branched, sometimes 
several times; herbage minutely puberulent, sometimes slightly glandular; 
principal leaf blades (1.5) 2-3 (5) em. long, and (1.5) 2-2.5 (3.5) em. wide, 
usually broadly ovate; leaf margins entire to sinuately or repandly few- 
toothed to dentate; petioles two-thirds the length of the blade to equalling it; 
corolla yellow, sometimes becoming bluish in age, or when dried and pressed, 
10-15 mm. long, its limb reflexed when fully open; anthers yellow, 2.5-3 mm. 
long; filaments having a few long hairs growing on them; calyx at anthesis 
usually 3-6 mm. long on peduncles 5-10 times their length; fruiting calyx 
usually 2-3 em. long and 1.5-2 em. wide. 

12a. P. crassifolia Bentham, var. crassifolia. P. cardiophylla Torrey, 
Bot. Mex. Bound. 153. 1859; P. crassifolia var. cardiophylla (Torr.) 
Gray, Synoptic Flora 2(1): 235. 1878; P. muriculata Greene, Bull. Calif. 
Acad. 1: 209. 1885. 

Leaves thick, entire to sinuately or repandly few-toothed; flowering 
calyx usually 4-6 mm. long on peduncles 6-7 times their length; corollas 
yellow, sometimes with brownish centers. 

TYPE: Bay of Magdalena, Lower California, Mexico; not seen. 

Variety crassifolia grows on deserts, canyon floors, rocky hillsides, and 
mountains, principally in Arizona, and California; it flowers from March 
through October. 

Selected from 185 sheets of 157 collections: AgizoNA: Cochise Co.: Blumer 
90 (ariz); Mohave Co.: Harrison, Kearney and Fulton 7549 (ariz); Pima Co.: 
Harrison and Kearney 7238 (xv); Pinal Co.: Gillespie 8919 (Ny, vc); Yavapai 


160 Rhodora [ Vor. 60 


Co.: Peebles, Harrison and Kearney 7431 (Ny); Yuma Co.: Benson 1080? 
(ARIZ); CALIFORNIA: Imperial Co.: Rose 36830 (oKL, mica); Inyo Co.: Clokey 
and Templeton 5776 (Ny, vc); Kern Co.: Munz, Johnston and Harwood 4034 
(ny); Riverside Co.: Clokey 6881 (Ny, uc); Mason 4185 (au, uc), Rose 36001 
(GH, vc); San Bernardino Co.: Munz 11720 (ariz, COLO, NY); San Diego Co.: 
Abrams 3160 (Gn, NY); Nevapa: Clark Co.: Clokey 8577 (coro, NY, uc); 
Clover 8235 (mica); Lincoln Co.: Kennedy and Gooding 10 (ARIZ, NY, UC). 
12b. P. crassifolia Bentham, var. versicolor (Rydberg) Waterfall, 
comb. et stat. nov., based on P. versicolor Rydb. Bull. Torr. Bot. Club 
22:307. 1895; P. genucaulis Aven Nelson, Bot. Gaz. 47: 430. 1909. 

Leaves thinner, usually dentate, but sometimes nearly entire; calyx 
usually 3-4 mm. long on peduncles 5-10 times their length; corolla yellow, 
usually some, or all, of them turning bluish in drying. 

TYPE: Rydberg selected no type, therefore Edward Palmer 622, col- 
lected at Guaymas, Mexico in 1887 is selected as the LECTOTYPE (NY); 
isolectotype (GH). 

Its habitat, distribution and flowering time are apparently similar to 
the above, but it is not so widespread. 

Selected from 71 sheets of 55 collections: Arizona: Gila Co.: King and 
Belden 2439 (arız); Mohave Co.: (?): Clover 6009 (ariz); Pima Co.: Toumey 
June 1, 1896 (GH, NY), Gould and Macbride 4128 (ariz, Gu, NY, UC); Pinal Co.: 
Thornber 5517 (arız, NY); Yuma Co.: Parker, Parker, Wright and Lowe 7816 
(COLO, NY, UC); CALIFORNIA: Imperial Co.: Wiggins 9606 (au, vc); Riverside 
Co.: Wiggins 9673 (Gu, Ny, Uc); Nevada: Clarke Co.: Train 1366 (Ny, vc). 

13. Physalis ixocarpa Brotero ex Hornemann, Hortus Regius Botanicus 
Hafniensis, Supplement 26. 1819; P. aequata Jacq. f. ex Nees, Linnaea 
6: 470. 1831. 

Annual, 15-60 cm. tall, branched, glabrous to rather sparsely vestite with 
short appressed hairs; leaf blades 2-7 cm. long, ovate to ovate-lanceolate; 
margins of the leaves dentate to sinuate-dentate to entire, on petioles about 
one-half as long as the blade to equalling it in length; corolla 7-15 mm. long, 
with 5 bluish-tinged dark spots on its limb which is recurved when fully open; 
anthers blue, about 3 mm. long, strongly twisted after dehiscence; flowering 
peduncles 3-5 mm. long; fruiting calyx usually 2-2.5 (3) em. long, nearly 
globose, often well-filled with the fruit; fruiting peduncles usually 3-8 mm. 
long. 

Fruiting material may often resemble P. virginiana var. subglabrata. 
It can be distinguished by its shorter peduncles. 

TYPE: None was selected by Hornemann, and no material was cited. 
Presumably a neotype should be selected, probably from Mexican collec- 
tions since the species seems to be native there. However the author 
prefers to defer this action until a more detailed study of the species from 
that area may be accomplished. The concept of the species is based upon 
the material cited later, which seems to be conspecific with Mexican ma- 
terial seen. 

This species is cultivated and escapes; it flowers through much of its 
growing season, 

Selected from 95 sheets of 81 collections: CANADA: Ottawa, Marie-Victorin, 
et al. 43923 (an); UNITED STATES: Carrronxia: Butte Co.: Yates 6127 


1958] Waterfall,—Genus Physalis in N. America 161 


(vc); Fresno Co.: Bacigalupi, Ferris and Wiggins 2491 (aH, NY, vc); Los 
Angeles Co.: Fosberg 53036 (Gn, xv); Riverside Co.: Conger Oct. 1909 (vc); 
San Bernardino Co.: Parish Sept. 1888 (vc); San Luis Obispo Co.: Miossi 
Aug. 5, 1840 (vc); Santa Barbara Co.: Bingham 29 (Nv); Ventura Co.: Pollard 
Oct. 27, 1945 (cono); DELAwARE: New Castle Co.: Commons Nov. 2, 1898 
(au); Sussex Co.: Churchill Sept. 11, 1908; Illinois: Adams Co.: Seymour Aug. 
1878 (DUKE); DuPage Co.: Moffett 3197 (Gu, oxra); Fulton Co.: Vasey 1862 
(aH); ManvLANp: McVaugh 134543, cult., originally from Mexico (wicH); 
MassAcnuusETTS: Middlesex Co.: Deane Sept. 24, 1884 (NEBC); Norfolk Co.: 
Fernald Sept. 26, 1908 (au); MrcuiGAN: Emmet Co.: Hoover 1943, seeds from 
the Orange Free State (DUKE, TEX, vc); Minnesota: Herb. Canby Sept. 1868 
(NY); New Jersey: Hunterdon Co.: Dodge July 18, 1899 (arcu); New Mexico: 
Rio Arriba Co.: Wooton 2697 (NY); Santa Fe Co.: Fendler 680 (au); New York: 
Ontario Co.: coll. unknown Aug. 2, 1887, raised from seeds from Palmer from 
Mexico (GH); Tompkins Co.: Hoisington 340, cultivated (OKLA); OREGON: 
Multnomah Co.: Nelson 3325 (Gu); PENNSYLVANIA: Fretz 1881 (uc); Phila- 
delphia Co.: Parker Sept. 9, 1874 (ny); Texas: Bexar Co.: Jermy 1904 (NY); 
Brewster Co.: Marsh 163 (Gn); Crockett Co.: Cory 29703 (au); Refugio Co.: 
Tharp Sept. 7, 1929 (micu); Webb Co.: Mackenzie 85 (NY); VERMONT: Chitten- 
den Co.: Flynn 4 (GH); Virainta: Clarke Co.: Young 485, raised in experi- 
mental plots (TEX); WASHINGTON: Klickitat Co.: Suksdorf 2284 (au, vc); 
Washington D. C.: Steele Sept. 20, 1899 (puKE); West VrnGiNIA: Rawleigh 
Co.: Tosh 650 (uc). 

14. Physalis Wrightii Gray, Proc. Amer. Acad. Arts and Sciences 10: 
63. 18795. 

Annual 30-90 em. tall, nearly glabrous, the few hairs short, stiff and ap- 
pressed; leaf blades ovate-lanceolate to linear-laneeolate, the principal ones 
usually 4-12 centimeters long on petioles 1.5-7 em. long; leaf margins usually 
irregularly and often coarsely dentate, sometimes regularly and saliently 
dentate; corolla a light yellow color, sometimes with a greenish tinge, rotate 
with very little tub», 15-23 mm. wide when fully open, with five hairy pads 
on its limb near the base, alternating with the stamens; anthers (2.8)3 (3.8) 
mm. long, yellow with a blue or blue-green tinge; filaments slender, somewhat 
exceeding the anthers in length; flowering calyx usually 4-5 mm. long on 
peduncles 5-12 times its length; fruiting calyx usually 2-2.5 cm. long and 
1.7-2 em. wide, on peduncles usually 2.5-6 cm. long, sometimes nearly filled 
by the fruit. 

TYPE: Charles Wright 1602, prairies along the San Pedro River, south- 
western Texas, 1851-52 (Gu); isotype (Ny); no other collections have been 
seen from Texas. 

Habitat, distribution and flowering time: Growing in deserts and moun- 
tains, but particularly in fields and other disturbed habitats, primarily in 
Arizona and California; flowering from July to November. 


Selected from 46 sheets of collections: ARIZONA: Cochise Co.: Griffiths 1579 
(ARIZ, NY); Gila Co.: Collom Sept. 15, 1934 (micu); Graham Co.: Richardson 
437 (ARIZ); Maricopa Co.: Wiggins 3860 (mich); Navajo Co.: Zuch 49 (ARIZ); 
Pima Co.: Pringle Aug. 1, 1894 (an, NY); Gould 3938 (ARIZ, TEX, vc); Pinal 
Co.: Arnold and Darrow Sept. 13, 1936 (au, uc); Yuma Co.: Thornber Sept. 
24, 1912 (arız); CALIFORNIA: Imperial Co.: Munz 11523 (ny); Kern Co.: 
Yates 6839 (vc); Los Angeles Co.: Wheeler 964 (vc); San Diego Co.: Brandegee 
July 1895 (uc); Texas: Wright 1602. (GH, NY). 


162 Rhodora [ Vor. 60 


15. Physalis angulata Linnaeus, Species Plantarum 1: 183. 1753; 
other synonymy under the varieties. 


Annual, 15-90 em. tall, glabrous, or with a few short appressed hairs es- 
pecially on the younger parts; blades of principal leaves usually 4-10 em. long, 
ovate to lanceolate, or sometimes linear-lanceolate; margins of the leaves ir- 
regularly and sometimes coarsely or saliently toothed, or entire, on petioles 
1-4 em. long; corolla yellowish, not dark spotted, usually 4-10 mm. long; 
anthers usually 1-2.3 mm. long, bluish, on slender filaments; flowering calyx 
usually 3-5 mm. long with calyx lobes 1-2.5 mm. long; flowering peduncles 
5-40 mm. long; fruiting calyx usually 2-3 em. long and 1.5-2.5 em. wide on 
peduncles 10-40 mm. long. 


15a. P. angulata L., var. angulata. P. Linkiana Nees, Linnaea 6: 
471-472. 1831; P. angulata L., var. Linkiana (Nees) Gray, Proc. Amer. 
Acad. Arts and Sciences 10: 64. 1875. 

Leaves ovate to ovate-lanceolate; corolla usually 6-10 mm. long; flower- 
ing calyx usually 4-5 mm. long with calyx lobes 2-2.5 mm. long; flowering 
peduncles usually 5-15 mm. long; fruiting calyx on peduncles usually 20-30 
mm. long, shorter than to equalling the length of the fruiting calyx. 

Variety angulata grows in open woods, pastures, ditches, fields, and 
various disturbed habitats in the extreme eastern states, and in the south- 
'astern states as far west as eastern Oklahoma and Texas; it flowers from 
May to September. 


Selected from 104 sheets of 77 collections: ALABAMA: Coosa Co.: Pollard 
and Ball 263 (Gn, Ny); Lee Co.: Earle and Baker Aug. 11, 1897 (Ny); Tusca- 
loosa Co.: Pollard and Mazon 330 (mich, NY); ARKANSAS: Arkansas Co.: 
Moore 82748 (OKLA, UARK); Ashley Co.: Demaree 16362 (Ny); Chicot Co.: 
Demaree 18567 (Ariz, NY); Hempstead Co.: Buckholz 388 (UARK); Jefferson Co.: 
Demaree 1398? (Ny); Pulaski Co.: Merrill 672 (UARK); Connecticut: Hartford 
Co.: Bissell Aug. 19, 1904 (GH); DELAWARE: Commons Aug. 15, 1877; FLORIDA: 
Collier Co.: Moldenke 5761 (xy); Columbia Co.: Straub 36 (GH); Duval Co.: 
Curtiss 5737 (Gu, NY, Uc); Gadsden Co.: Berg (Ny); Gulf Co.: Chapman 4345 
(au, NY); Hillsborough Co.: Deam 2746 (au); Lake Co.: Nash 1052 (au, 
MICH, uc); Lee Co.: Hitchcock 238 (Gu, Ny); Leon Co.: Godfrey 52385 (DUKE); 
Pinellas Co.: Deam 2905 (au); Polk Co.: McFarlin 6569 (mica); Volusia Co.: 
Hood 9 (an); Grorata: Decatur Co.: Thorne 4629 (an); De Kalb Co.: Small 
Sept. 11, 1894 (ny); McDuffie Co.: Bartlett 1662 (micu); Louisiana: Baton 
Rouge Parish: Chamblis 17 (ny); Correll 9567 (DUKE, GH, NY); Tangipahoa 
Parish: Correll 9240 (pu KE); MississiPrr: Harrison Co.: Tracy 6476 (Gau, NY); 
Jackson Co.: Pollard 1123 (Gu, Ny); New Jersey: Brunswick Co.: Godfrey 10085 
(TEX); Camden Co.: Beringer Sept. 1891 (Micu); Norra CAROLINA: Moore 
Co.: Oosting 34777 (Duke, PH); New Hanover Co.: Williamson Sept. 1, 1900 
(Ny, PH); Wilson Co.: Randolph and Randolph July 7, 1922 (Gu); OKLAHOMA: 
Delaware Co.: Wallis 2728 (okLA); Oklahoma Co.: Waterfall 2349 (oKL); 
Pottawatomie Co.: Barkley 395 (oki); Sours Carona: Charleston Co.: 
Moldenke 5196 (ny); in cultis Curtis (au); Texas: Angelina Co.: Cory 10665 
(an); Brazos Co.: Parks Dec. 1, 1946 (rex); Gonzales Co.: Tharp Nov. 23, 
1935 (mıca); Harris Co.: Boon 481 (rex); Houston Co.: Cory 26121 (an); 
Jefferson Co.: Tharp Sept. 9, 1937 (rex); Wood Co.: Cory 57671 (coro); 
VırGINIA: Princess Anne Co.: Fernald and Long 10881 (au); Southampton 
Co.: Fernald and Long 13742 (an). 


1958] Waterfall,—Genus Physalis in N. America 163 


15b. P. angulata L., var. pendula (Rydberg) Waterfall, comb. et stat, 
nov., based on P. pendula Rydb. in Small, Flora of the Southeastern 
United States 983. 1903. 


Similar to var. angulata, but leaves sometimes narrower; flowering calyx 
usually 3 mm. long, sometimes 4 mm. long; with calyx lobes about 1 mm. long; 
flowering peduncles usually 15-40 mm. long; fruiting peduncles usually 20—40 
mm. long, equalling the fruiting calyx (which is usually 20-25 mm. long) to 
three times its length. 


TYPE: In describing this species, Rydberg did not select a type, stating 
that it was the taxon that he had originally called P. lanceifolia, or at least 
the part of it occurring from Illinois to Texas. He cited several collections 
in his treatment of the genus (1896), and from among these F. L. Harvey 65 
“central and southern Arkansas" (UARK) is selected as LECTOTYPE. 

This variety grows in river valleys, bottom woods, fields and various 
disturbed sites, primarily in Oklahoma and Texas, but extending north to 
Illinois; it flowers from June through September. 


Selected from 125 sheets of 104 collections: ARKANSAS: Conway Co.: Moore 
420449 (UARK); Garland Co.: Demaree 20471 (Ny, vc); Harvey 65 (micH); 
Little River Co.: Moore 510682 (uark); Prairie Co.: Demaree 15498 (NY); 
Pulaski Co.: Merill 725 (uark); InuiNOoms: Alexander Co.: Palmer 16628 
(pH); Cook Co.: Umbach Aug. 3, 1897 (mich, Ny, PH); St. Claire Co.: Eggert 
Sept. 16, 1893 (GH); Union Co.: Vasey 1862 (au); Vasey (Ny); Kansas: 
Douglas Co.: McGregor 607 (KANU); Geary Co.: Hitchcock 775 (GH, NY); 
Linn Co.: Rydberg and Imler 77 (Ny); Miami Co.: McGregor 11048 (KANU); 
Sedgwick Co.: Horr 6191.1 (KANU); Louisiana: Hale (GH); MASSACHUSETTS: 
Middlesex Co.: Perkins Oct. 22, 1880 (Ny); Missounr: Jackson Co.: Mackenzie 
May 10, 1896 (Ny); Jasper Co.: Palmer 3093 (Ny); St. Louis Co.: Eggert Sept. 
7, 1887 (pH, vc); Engelmann 324 (GH); OKLAHOMA: Alfalfa Co.: Waterfall 9970 
(OKLA); Blaine Co.: Waterfall 2387 (oKL, vc); Cherokee Co.: Wallis 1468, 
1891 (OKLA); Creek Co.: Bush 397 (an); Custer Co.: Palmer 12555 (TEX, vc); 
Garvin Co.: Andrews 133 (oki); Logan Co.: Smith 889 (oki); Murray Co.: 
Robbins 2729 (oki); Muskogee Co.: Little 188 (oKL); Oklahoma Co.: Waterfall 
2091 (OKLA, GH); Osage Co.: Stevens 2115 (GH, NY, OKL, OKLA); Payne Co.: 
Coryell 388, 596 (okrA); Pittsburg Co.: McClary 66 (ok); Pontotoc Co.: 
McCoy 852, 1267, 1915 (okrA); Pottawatomie Co.: Van Vleet July 12, 1905 
(okL); Texas: Bexar Co.: Metz 64 (ny, uc); Bowie Co.: Plank May 9, 1891 
(NY); Brazos Co.: Reeves 62 (Gu); Calhoun Co.: Gentry 49 (TEX, LIL); Colorado 
Co.: Bush 333 (Gu, NY); Dallas Co.: Hall 504 (an, Ny); DeWitt Co.: Riedel 
Aug. 3, 1941 (rex); Harris Co.: Boon June 22, 1943 (TEx); Jackson Co.: 
Tharp Aug. 8, 1941 (tex); Lamar Co.: Strandtman 10 (TEX); McLennan Co.: 
York 46232 (okr, TEX); Refugio Co.: Tharp Dec. 4, 1928 (TEX); San Patricio 
Co.: Cory 45389 (au); Tarrant Co.: Ruth 1242 (Ny); Titus Co.: Jones 10 
(TEX); Travis Co.: Tharp 1717 (rex); Washington Co.: Brackett July 15, 1938 
(GH, TEX). 

15e. P. angulata L., var. lanceifolia (Nees) Waterfall, comb. et stat. 
nov., based on P. lanceifolia Nees, Linnaea 6: 473. 1831. 

Similar to var. pendula, but leaves lanceolate to linear-lanceolate, and 
corolla usually only 4-5 mm. long; anthers often only 1-1.5 mm. long. 

TYPE: No collections were cited by Nees who said “Habitat in Peruvia 


164 Rhodora [ Vor. 60 


(Ruiz et Pavon); in Mexico (Herb. Hort. Reg. Ber.)". A Neotype should 
be selected, but preferably after a study of Peruvian and Mexican material. 
Specimens cited below appear to be similar to a number of Mexican collec- 
tions seen by the author. 

Variety lanceifolia grows in wet areas, river valleys, fields and other 
disturbed habitats in California, Arizona, New Mexico and to a lesser 
extent in Texas and southern Oklahoma, often being atypical in the latter 
two states, probably due to gene interehange with var. pendula; a few 
Florida collections are also referred here; it flowers usually in June to 
September, or as late as November in the warmer parts of its range. 

Selected from 64 sheets of 41 collections: AnrzoNaA: Cochise Co.: Thornber 
2627 (au), put here because the anthers are only about 1 mm. long, even though 
the leaves are broad; Pima Co.: Pringle Aug. 5, and Sept. 5, 1884 (xv, PH, vc); 
Pinal Co.: Kearney 15067 (ariz); Yuma Co.: Schott 2 (ny); CALIFORNIA: 
Butte Co.: Heller 13355 (au); Fresno Co.: Bacigalupi, Ferris and Wiggins 2488 
(GH, NY, Uc); Imperial Co.: Parish 8337 (au); Los Angeles Co.: Wheeler 965 
(uc); Merced Co.: Hoover 1599 (uc); San Diego Co.: Spencer 1014 (am); 
Stanislaus Co.: Hoover 2442 (uc); Tulare Co.: Michener and Bioletti 1893 
(NY); FLortpa: Dade Co.: Small and Carter 649 (Ny, pH); Kevy Co.: Small, 
Small and DeWinkeler 10036 (Ny); Monroe Co.: Eyles 8213 (an, oki); New 
Mexico: Dona Ana Co.: Archer 489 (micu); Kearney and Peebles 15073 
(ARIZ); OKLAHOMA: McCurtain Co.: Waterfall 7604 (oKL, OKLA); Oklahoma 
Co.: Waterfall 2893, near var. pendula (oKL); Payne Co.: James 37 (OKLA); 
Texas: Brewster Co.: Cory 31275 (au); Cameron Co.: Runyon 4243 (TEX); 
Llano Co.: Bray 10 (Ny); Presidio Co.: Warnock T 164 (au, TEX); Refugio Co.: 
Tharp Sept. 7, 1929 (rex); Travis Co.: Tharp, Warnock and Barkley Nov. 31, 
1945, atypical material, perhaps intermediate with var. pendula (COLO, DUKE, 
GH, NY, OKL, OKLA, UARK, UC). 

16. Physalis pubescens Linnaeus, Species Plantarum 1: 183. 1753. 
Synonymy listed under the varieties. 

Plants annual, 15-60 cm. tall, villous or viscid-villous vestite, sometimes 
with granular glands, sometimes glabrate; blades of principal leaves usually 
3-10 em. long, narrowly to broadly ovate, on petioles half as long to about 
equal them in length; margins of the leaves toothed to entire; corolla 6-10 
mm. long, yellow with 5 prominent dark spots on the limb near its base; 
anthers 1.5-2 (2.4) mm. long, blue; flowering calyx 4-7 mm. long, its lobes 
2-4 mm. long; flowering peduncles 3-12 mm. long; fruiting calyx 2-4 em. long 
and 1.5-2.5 em. wide, 5 angled, on peduncles 5-20 mm. long. 

16a. P. pubescens L., var. pubescens. P. turbinata Medicus, 
Academia Theodora-palatina 4: 188-192. 1780; P. barbadensis Jacquin, 
Miscellanea Austriaca Sive Plantarum Selectarum 360. 1781; P. obscura, 
var. viscido-pubescens Michx., Flora Boreali-Americana 1: 149. 1803; 
Alicabon barbadense (Jacq.) Rafinesque, Sylva Telluriana 56-57. 1838; 
P. viscido-pubescens (Michx.) Dunal, in DeCandolle's Prodromus 13(1): 
442. 1852; P. floridana Rydberg, in Small, Flora of the Southeastern 
United States 983. 1903. 

Plants more or less villous; leaf blades usually with 5-8 teeth on each 
side, usually not translucent; flowering peduncles 3-7 mm. long; fruiting 
ealyees 2-3 em. long on peduncles usually 5-9 mm. long. 


1958] Waterfall, —Genus Physalis in N. America 165 

TYPE: “In India utraque.” A photograph of the type is in the Arnold 
Arboretum's collection of photographs of specimens in the Linnaean 
Herbarium, London. 

Variety pubescens grows in swamps, margins of lakes, sand dunes, brush, 
fields and other disturbed habitats, primarily from Florida to Texas in 
our area; it flowers from May through November. It is a widespread 
pantropical taxon. 

Selected from 81 sheets of collections: FLonipA: Broward Co.: Moldenke 480 
(NY); Dade Co.: Small and Small 4632 (Duxe, Ny); Franklin Co.: Chapman 
3055b (an, NY); Highlands Co.: Moldenke 5417 (Ny); Hillsboro Co.: Churchill 
March 28, 1936 (GH); Lake Co.: Nash 1251 (Gu, MICH, NY, vc, PH); Lee Co.: 
Tracy 7612 (au, NY); Monroe Co.: Killip 41456 (Ny, uc); Pinellas Co. : Williams 
Mar. 12, 1926 (puKE); Polk Co.: McFarlin 5924 (mica); St. Lucie Co.: Small 
8507 (GH, NY); ILLiNois: Jackson Co.: Vasey (au); Louisiana: Hale (aH); 
Texas: Bexar Co.: Metz 771 (micu); Brazos Co.: Moncreif 1476 (TEX); 
Gonzales Co.: Tharp Aug. 12, 1940 (rex); Harris Co.: Fisher Oct. 9, 1917 
(uc); Hidalgo Co.: Walker 8 (Gu, TEx); Jackson Co.: Warnock 105 (TEX); 
Jefferson Co.: Tharp Sept. 10, 1937 (au, TEX); McLennan Co.: Smith 59 
(rex); Newton Co.: Tharp 52141 (au); Nueces Co.: Tharp and Brown 48-165 
(TEX); Travis Co.: Tharp Nov. 8, 1929 (Gu, oxra, TEX); Willacy Co.: Johnston 
542221 (TEX); Williamson Co.: Wolcott 314 (TEx). 

16b. P. pubescens L., var. glabra (Michx.) Waterfall, comb. nov., 
based on P. obscura Michx., var. glabra Michx. Flora Boreali-Americana 
1: 149. 1803. P. obscura Michx., le. P. hirsuta Dunal, var. repando- 
dentata Dunal, in DeCandolle's Prodromus 13(1):445. 1852; P. barbaden- 
sis Jacq., var. obscura (Michx.) Rydb., Mem. Torr. Bot. Club 4: 327. 
1896; P. barbadensis Jacq., var. glabra (Michx.) Fernald, Ruopora 51: 
82. 1949. 

When Michaux described P. obscura he immediately divided it into two 
varieties, var. glabra and var. viscido-pubescens. The present author in- 
terprets var. glabra as being the “typical” variety. 

This material has been referred recently, by some American authors, to 
P. turbinata Medicus, Academia Theodora-palatina 4: 188-192. 1780. 
However, in describing this species Medicus says “Die Hauptstamme und 
nebenaste sind vierkandigt, haarich und rotlich violet-braun." One might 
dismiss the reddish violet-brown color, as either not necessarily being a 
characteristic of the whole taxon, or as possibly not being retained in 
herbarium specimens, but it seems dubious if the term “hairy” would be 
used to describe nearly glabrous, or slightly puberulent specimens. It 
seems that the taxon described by Medicus, at least as the name has been 
applied in our flora, is more likely referable to var. pubescens. 

Plants glabrous or sparingly puberulent, but not villous as in the other 
varieties; blades of the principal leaves usually 2-7 em. long, ovate, often 
rather broadly so, acuminate in many specimens; margins of the leaves ir- 
regularly toothed, sometimes saliently so; petioles about equalling the blades 
in length; anthers 1.8-2.4 mm. long, bluish; flowering calyx 5-7 mm. long 
with narrow lanceolate-acuminate lobes 2.5-4 mm. long; flowering peduncles 
5-12 mm. long; fruiting calyx 3-4 em. long, ovate or broader in outline, often 
acuminate at the apex, on peduncles 1-2 em. long. 


166 Rhodora [ Vor. 60 


TYPE: In the Herbarium of Michaux, Mus. Hist. Nat., Paris; photograph 
in the Gray Herbarium. 

Variety glabra grows in open woods, creek sides, valleys, vards and other 
disturbed habitats, mostly in southeastern coastal states from North 
Carolina to Texas, and inland in Arkansas and Missouri; it usually flowers 
from July through October. 

Selected from 47 sheets of 34 collections: ALABAMA: Crenshaw Co.: Reed 2105 
(rex); Mobile Co.: Mohr Aug. 1883 (mic); ARKANSAS: Drew Co.: Demaree 
16498 (NY); Fulton Co.: Bush 961 (Ny); Hot Springs Co.: Demaree 19471 
(NY); Logan Co.: Palmer 24209 (UARK); Saline Co.: Moore 53-311 (UARK); 
FLoripa: Dade Co.: Tatnell 620 (vun); Gadsden Co.: Curtiss 5896 (GH, UC); 
GEoRGIA: Calhoun Co.: Thorne 7338 (Gu); CALIFORNIA: San Diego Co.: 
Jones March 1882 (pH); Louisiana: Calcasieu Parish: Correll and Correll 9566 
(DUKE, GH, NY, PH); Natchitoches Parish: Palmer 8777 (vu); Vermilion Parish: 
Tharp July 27, 1929 (rex); Mississippi: Oktibbeha Co.: Pollard 1338 (Gu); 
Missourt: Barry Co.: Bush 547 (Ny); Butler Co.: Eggert July 1893 (xy, vc); 
Madrid Co.: Bush 189 (Gu, NY); NORTH CAROLINA: Curtis (GH); PENNSYLVANIA: 
Bucks Co.: Moyer (rn); Texas: Bowie Co.: Heller and Heller 4253 (Gu, NY, 
pH); Harris Co.: Hall 503 (au, ny); Newton Co.: Tharp 42-141 (Gu, TEX); 
Orange Co.: Tharp 2518 (rex); Rusk Co.: Reverchon 3239 (NY). 

16c. P. pubescens L., var. integrifolia (Dunal) Waterfall, comb. nov., 
based on P. hirsuta Dunal, var. integrifolia Dunal, in DeCandolle, Prodro- 
mus 13(1): 445. 1852. 

Plants more or less villous; leaf blades often entire, sometimes 3-4 (rarely 
more) more or less prominent teeth on each side, translucent or semitrans- 
parent; fruiting calyx 2-3 em. long on peduncles 5-9 mm. long. 

Included here are the plants, primarily of the northeastern United 
States, which have been referred to P. pubescens by recent American 
authors. 

TYPE: “Physalis, n. 30, un. itin., Frank e sylvaticis agri Cincinnati civ. 
Ohio," presumably in the De Candolle collection in the Conservatoire et 
Jardin Botaniques, Genève, Switzerland. 

Variety integrifolia grows on creek banks, lake shores, woods, hills and 
various disturbed habitats, mostly from Pennsylvania to Iowa and south to 
Florida and south central Texas, but also in southern New Mexico, Arizona 
and California; it is less frequent in southeastern United States than var. 
pubescens. 

Selected from 166 sheets of 132 colleetions: ALABAMA: Jefferson Co.: Karle 
June 18, 1899 (xv); Tuscaloosa Co.: Pollard and Mazon 331 (GH); ARIZONA: 
Pima Co.: Toumey Aug. 30, 1895 (uc); ARKANSAS: Baxter Co.: Moore 610540 
(OKLA, UARK)} Clay Co.: Demaree 20311 (OKLA, NY, UC); Crawford Co.: 
Demaree 15300 (xy); Garland Co.: Demaree 16190, 21841, 20422 (NY); Newton 
Co.: Moore 430237 (OKLA, UARK); Perry Co.: Demaree 20168 (Ny); Polk Co.: 
Moore and Williams Aug. 15, 1951 (vark); Pulaski Co.: Demaree 16640 
(ny); Sevier Co.: Demaree 9913 (NY, vc); Union Co.: French 500150 (UARK); 
Washington Co.: Giles 429 (UARK); Yell Co.: Demaree 20109 (NY); CALIFORNIA: 
Colusa Co.: Stinchfield 460 (xv); Imperial Co.: Thomas (Gu, xv); Lake Co.: 
Baker 11226 (vc); San Diego Co.: Orcutt Mar. 6, 1883 (wticu); Tulare Co.: 
Congdon Oct. 8, 1881 (vc); FLortpa: Dade Co.: Small and Moiser 5902 (GĦ, 


1958] Waterfall, —Genus Physalis in N. America 167 


ny); Leon Co.: Godfrey 52473 (puKE); ILLINOIS: Adams Co.: Seymour Sept. 
26, 1876 (DUKE); Massae Co.: Gleason 2630 (GH); INDIANA: Grant Co.: Deam 
15287 (NY); Lawrence Co.: Kriebel 2564 (pUKE); Putman Co.: Banker 1499 
(NY); Tippecanoe Co.: Boot Oct. 6, 1895 (GH); Whitely Co.: Friesener 16539 
(Gu, NY); Kansas: Riley Co.: Norton 366 (au, Ny); KENTUCKY: Bell Co.: 
Lloyd Aug. 10, 1888 (v); Hickman Co.: McFarland and Anderson 2223 
(NY); ManYLAND: Montgomery Co.: Blanchard Aug. 12, 1892 (Ny); Worcester 
Co.: Canby Sept. 1863 (xy); Massacuuserts: Suffolk Co.: Perkins Sept. 6, 
1881 (NEBC); Missouri: Butler Co.: Eggert July 1893 (uc) on sheet with var. 
obscura; Jackson Co.: Bush 6423 (Gu, NY); Jasper Co.: Demaree 4424 (OKLA, 
UARK); Moniteau Co.: Steyermark 70814 (UARK); Newton Co.: Palmer 32492 
(NY); Osage Co.: Jeffrey 366 (GH); Phelps Co.: Kellogg 196 (NY, TEX, UC); St. 
Louis Co.: Eggert Aug. 14, 1891 (rex, uc); New Mexico: Rusby 310, Burro Mts. 
(au, NY); Nortu Carouina: Brunswick Co.: Blomquist 4811 (bUKE); Carteret 
Co.: Lewis 234 (Ny); Washington Co.: Correll 1921 (puKE); Onto: Moldenke 
13543 (OKLA); Franklin Co.: Gleason Sept. 5, 1904 (GH); Hamilton Co.: Lloyd 
2209 (MicH); Lake Co.: Werner 141 (GH); OKLAHOMA: Cherokee Co.: Waterfall 
9661 (OKLA); Delaware Co.: Wallis 2732 (oKLA); Johnston Co.: Houghton 357214 
(NY); Murray Co.: Hopkins and Cross 6429 (oKL); Muskogee Co.: Waterfall 
10139 (OKLA); Ottawa Co.: Stevens 2530 (Gu, Ny); Payne Co.: Thompson 82 
(OKLA); PENNSYLVANIA: Allegheny Co.: Porter Aug. 28, 1896 (GH, NY); SouTH 
CanoLINA: Berkeley Co.: Godfrey and Tryon 622 (GH, NY); TENNESSEE: 
Cheatham Co.: Svenson 10395 (uc); Davidson Co.: Svenson 9494 (GH); Hamil- 
ton Co.: Clalmgh 101 (puxe); Texas: Cameron Co.: Johnston 542210 (TEX); 
Dallas Co.: Reverchon 382 (Gu); Gonzales Co.: Tharp 51-467 (OKLA, TEX); 
Jackson Co.: Warnock 105 (Ny); Travis Co.: Armor 5508 (OKLA) approaching 
var. pubescens; Willacy Co.: Davis and Johnston 53256.15 (TEX); VIRGINIA: 
Henrico Co.: Fernald and Long 12794 (Gu): Isle of Wight Co.: Fernald and Long 
13442 (Gu); James City Co.: Fernald and Long 13441 (an); Loudon Co.: Holms 
Aug. 1888 (aniz, Ny); Nansemond Co.: Fernald and Long 10810 (Gu); North- 
ampton Co.: Canby Sept. 1878 (Ny); Page Co : Steele and Steele 197 (Gu, NY); 
Princess Anne Co.: Fernald and Long 4167, 4168, 10809 (au); WEST VIRGINIA: 
Mertz Sept. 22, 1878 (NY). 

16d. P. pubescens L., var. grisea Waterfall, var. nov., Planta grisea, 
nune villosa nune brevipilosa, nune glandulari-farinacea; folis ovatis 
sinuato-dentatis; calycibus fructu a pendunculis 5-9 mm. longis. 

Stems densely covered with long, jointed hairs, or with long and short 
hairs mixed, or densely short viscid-hairy; leaves usually short hairy, some- 
times with granular glands, the surfaces having a greyish appearance; leaf 
blades ovate, coarsely and irregularly 6-9 dentate, or sinuate dentate nearly 
to their bases; fruiting calyces on peduncles 5-9 mm. long. 

This is the taxon, primarily of the northeastern United States, that has 
been passing as P. pruinosa L. However the photograph of the type of 
P. pruinosa in the Arnold Arboretum of Harvard University shows a plant 
with a more prominent acumination of the leaf blade, and a much longer 
flowering peduncle than is found in any of our material. The author has 
been unable to determine the application of the name, P. pruinosa. He 
has seen no material comparable with the photograph of the type. 

TYPE: Walter Deane Sept. 24, 1884, Cambridge, Mass. (an; isotype: 
NY). 


168 Rhodora [ VoL. 60 


Variety grisea grows on mountainsides, wooded slopes, roadsides, in 
gardens and various disturbed habitats, principally in northeastern United 
States; it usually flowers in August, September and October. 

Selected from 114 sheets of 102 collections: CANADA—Onvario: Macoun 
34524 (NY); UNITED STATES —AraABAMA: Baldwin Co.: Dukes 118 (NY); 
CALIFORNIA: Inyo Co.: Roos and Roos 6221 (uc); Connecticut: New Haven 
Co.: Blewitt 1381 (NEBC); Detawarr: New Castle Co.: Latnall 1882 (an); 
FromipaA: Chapman (Ny); GEoncia: DeKalb Co.: Small Aug. 1-6, 1895 (NY); 
Iuuinors: DuPage Co.: Umbach 12484 (au); Macon Co.: Mills Sept. 29, 1940; 
InpIANA: Lawrence Co.: Kriebel 2538 (Duke, GH); Kansas: Imler 68 (NY); 
Kentucky: Short 1840 (NY); Maine: Cumberland Co.: Chamberlain 1127 
(NEBC); MassaAcHUsETTS: Barnstable Co.: Collins 950 (NeBc); Bristol Co.: 
Hervey (NEBC); Dukes Co.: Bicknell 7704, 7706, 7693 (Ny); Essex Co.: Morong 
Aug. 1, 1868 (Ny); Hampshire Co.: Torrey and S.J.E. Sept. 16, 1943 (DUKE); 
Middlesex Co.: Fernald Sept. 26, 1908 (Gu); Nantucket Co.: Flynn July 30, 
1904 (NeBC); Norfolk Co.: Kidder Aug. 23, 1888 (NEBC); Plymouth Co.: 
Williams Aug. 21, 1898 (NEBC); Suffolk Co.: Young Sept. 1878 (GH, NEBC); 
Worcester Co.: Woodward 2 (Gu); MrcniGAN: St. Claire Co.: Dodge Aug. 25, 
1906; Missouri: Barry Co.: Bush 564, 469 (Ny); Christian Co.: Blankenship 
Aug. 1, 1895 (au); Jackson Co.: Mackenzie ? (micu); Jasper Co.: Bush 10402 
(Gu, NY); Taney Co.: Bush 170 (au); New Jersey: Hastings Sept. 6, 1917 
(NY) foot of Palisades; New York: Chemung Co.: Lucy 7825, 11098 (NY); 
Oswego Co.: Sheldon 6008 (uc); Tompkins Co.: Hoisington 312 (oki); Wash- 
ington Co.: Burnham Sept. 25, 1896 (Gu); North Carona: Granville Co.: 
Godfrey 2060 (au); Jackson Co.: Thazter June-July 1887 (Gn); Swain Co.: 
Beardslee and Kofoid Aug. 15, 1891 (Gn); OnEGoN: Tillamook Co.: Lloyd 
Sept. 10, 1894 (Ny); PENNSYLVANIA: Bucks Co.: Fretz Sept. 7, 1901 (an); 
Lancaster Co.: Small Sept. 1889 (GH); Westmoreland Co.: Shafer and Medayer 
182 (vc); Ruope Istanp: Providence Co.: Leland Sept. 18, 1881 (NEBC); 
TENNESSEE: Knox Co.: Ruth 3411 (ny); Texas: De Witt Co.: Riedel Aug. 3, 
1941 (TEX); VERMONT: Bennington Co.: Ames May 1885 (mica); Chittenden 
Co.: Flynn 3 (au); Rutland Co.: Eggleston 1510 (Gu, NEBC); VirGInra: Bedford 
Co.: Curtiss Oct. 3, 1871 (GH); Page Co.: Steele and Steele Aug. 28, 1901 (an, 
NY); WaAsHINGTON: Klickitat Co.: Suksdorf 2285 (xy); Yakima Co.: Hen- 
derson 2496 (Gn). 

17. P. foetens Poiret, var. neomexicana (Rydb.) Waterfall, comb. et 
stat. nov., based on P. neomexicana Rydb., Mem. Torr. Bot. Club 4: 325- 
320. 1895. 

Plants annual, 10-60 cm. tall, usually branched; indument short (0.5-1 
mm. long) and usually dense, more or less yellowish or brownish capitate- 
glandular; leaf blades 3-6 em. long, ovate to oblong-ovate or lanceolate- 
ovate, their margins toothed, or sometimes sinuate-toothed; petioles one-half 
to three-fourths as long as the blades; corollas 6-7 mm. long, bluish spotted; 
anthers (0.3) 1-1.5 (2) mm. long, bluish, on filiform filaments; flowering calyces 
3-4.5 mm. long, on peduncles usually 1.5-3 mm. long; fruiting calyces 2-3 
em. long, more or less ovate in outline, sharply 5-angled, on peduncles mostly 
4-7 mm. long. 

This variety differs from var. foetens of Mexico primarily in its shorter 
anther-length (2-3 mm. in var. foetens), usually shorter corolla (as much as 
1 em. long in some Mexican material) and in having fewer yellowish or 
brownish capitate glands than var. foetens. 


1958] Waterfall, —CGenus Physalis in N. America 169 


TYPE: In describing P. neomexicana, Rydberg cited several collections, 
but designated none of them as type. From among those cited, Fendler 
678 (an) is selected as the LECTOTYPE. A second sheet of the same collec- 
tion, an isolectotype, is in the same herbarium. 

Variety neomexicana grows in the mountains, often with junipers and 
pines, and in adjacent areas, including cultivated fields, in New Mexico 
and adjacent Colorado and Arizona; it flowers in June through October. 


Selected from 48 sheets of 40 collections: Arizona: Graham Co.: Bohrer 
409 (arız); Greenlee Co.: Gould and Haskell 4080 (vc); Maricopa Co.: Rusby 
310 (micu); Navajo Co.: Wooton September 13, 1913 (arız); Pima Co.: 
Toumey Aug. 30, 1894 (NY); Yavapai Co.: Wilcox Sept. 1918 (Ariz); COLORADO: 
Porter July 1872 (ru); El Paso Co.: Livingston 497 (DUKE); New Mexico: 
Jolfax Co.: Standley 13869 (ny); Bernalillo Co.: Ellis 287 (ny); Grant Co.: 
Rusby Oct. 1881 (mica); Lincoln Co.: Skehan 60 (au, NY); Wooton 633, 635 
(NY); Rio Arriba Co.: Parker and McClintock 6449 (ariz, vc); San Miguel 
Co.: Standley 4920 (au, Ny), Nelson 11568 (uc); Santa Fe Co.: Heller and 
Heller 3803 (au, wv); Sierra Co.: Metcalfe 1210 (aH, Ny, vc); Socorro Co.: 
Metcalfe 425 (ny); Torrance Co.: Parker and McClintock 6529 (NY); Socorro 
or Grant Co.: Rusby 309, Mogollon Mts. (mica). 


18. Physalis latiphysa Waterfall, sp. nov. Planta annua, 15-45 em. alta, 
ramosa, plus minusve villosa; foliorum laminis 5-7 em. longis, ovatis vel 
ovatis-rotundis, integris vel paucidentatis, acuminatis; corollis maculatis, 
4-6 mm. longis; antheris coeruleis, 1.5-2 mm. longis; calycibus fructu 
(2.5) 3-4 em. latis; pedunculis 1-1.5 em. longis. 


Annual, 15-45 em. high, branched, more or less villous; blades of the prin- 
cipal leaves 5-7 em. long, ovate to ovate-rotund, thin and translucent, their 
margins from entire to having a few teeth, acuminate; petioles 1.5-7 em. long; 
corollas yellow, dark-spotted, small, 4-6 mm. long; flowering calyces 3-4 mm. 
long with lobes about half that long, on peduncles 3-8 mm. long; fruiting 
calyces sparsely appressed-hairy, strongly 5-angled, 2.5-4 cm. long and (2.5) 
3-4 em. wide; fruiting peduncles 1-1.5 em. long; linear-subulate calyx lobes 
7-10 mm. long, extending 5-7 mm. beyond the body of the inflated fruiting 
calyx. 

TYPE: T. H. Kearney and R. H. Peebles 14425, Rondstadt Ranch, plain 
east of Baboquivari Mts., Pima Co., Arizona, Sept. 23, 1939. It is 
deposited in the Herbarium of the University of Arizona. 


In addition to the type, the following collections have been seen: ARIZONA: 
Pima Co.: Bartram 237 Santa Catalina Mts., east of Pima Canyon, Jan. 16, 
1920 (rm); Kearney and Peebles 10427, Toro Canyon, Baboquivari Mts., 
Sept. 30, 1934 (arız, MicH); Kearney and Peebles 14932, South Canyon, 
Baboquivari Mts., Aug. 31, 1940 (arız); Santa Cruz Co.: Harrison and Hope 
9058, Forty miles south of Tucson on Sasabe Road, Sept. 11, 1932 (ARIZ); 
Harrison and Fulton 8158, Nogales, Aug. 30, 1931 (arız); County undeter- 
mined; Harrison 9058, Robles to San Fernando, Sept. 10, 1932 (aH, MICH); 
Harrison, Kearney and Hope 8950 half-way from Sasabe to Robles, Aug. 21, 
1932 (arız); Kearney and Peebles 10576, Florida Canyon, Santa Rita Mts., 
Oct. 7, 1934 (ariz). 


19. Physalis missouriensis Mackenzie and Bush, Trans. Acad. Sci. 
St. Louis 12: 84-85. 1902. 


170 Rhodora [Vor. 60 


Annual, usually branched above, villous, sometimes with shorter hairs, 
often somewhat viscid; principal leaf blades 2-6 em. long, ovate to narrowly 
ovate, dentate to sinuately dentate, or sometimes entire; petioles one-third 
the length of, to nearly as long as the blades; corolla yellow, not dark spotted, 
7-10 mm. long; anthers (0.6) 1-1.2 mm. long, bluish, on slender filaments; 
flowering calyx 3-4 mm. long, its lobes 1-2 mm. long, on peduncles 3-6 mm. 
long; fruiting calyx 1.5-2.5 em. long, ovoid, on peduncles 5-10 mm. long. 

This species is easily distinguished from P. pubescens by its unspotted 
corollas and its smaller anthers. 

TYPE: K. K. Mackenzie 485, Rocky soil, Red Bridge, Jackson Co., 
Missouri. The type was deposited in the “Herbarium of K. K. Macken- 
ze."  Isotypes: GH, MICH, WIS. 

P. missouriensis grows in rocky woods and limestone barrens, mostly in 
Missouri, northeastern Kansas, western Arkansas, with two collections 
from adjacent Oklahoma, and one collection, dubiously referred here, 
from southwestern Texas; it flowers from June through October, 

Collections examined: ARKANSAS: Carroll Co.: Palmer 29310 (UARK); 
Hempstead Co.: Palmer 8955 (pu); Washington Co.: Moore 3008 (vARK); 
Moore and Iltis 430209 (OKLA, UARK); Giles 404 (uARK); J. T. B. 645 White 
Hiver (UARK); Kansas: Douglas Co.: Snow 2210 (Kanu); McGregor 9703 
(KANU); Marshall Co.: Horr 4610 (KANU); Riley Co.: Gates 18566 (Gu, TEX, 
Co.: Bush 162 (ok); Jackson Co.: Bush June 27, 1887 (au), Aug. 1888 (au, 
NY); 772 (au, NY), 4079 (au), 7334 (Gu), 7695 (Gu, NY), 12298 (Ny), 12298A 
(NY); Mackenzie 360 (ny), Aug. 23, 1896 (an, Ny); 485 (mic); Jefferson Co.: 
Prince July 4, 1883 (au); Phelps Co.: Kellogg Oct. 22, 1913 (rex); Platte Co.: 
Bush 11804 (Ny); St. Louis Co.: Eggert July 20, 1887 (au), Aug. 21, 1891 (NY); 
Pennell 11701 (vn); Taney Co.: Bush 173 (an, Ny); County undetermined; 
Blankenship 1893 (wy); Nelson 5 (Ny); OKLAHOMA: Muskogee Co.: Little 2568 
(OKL); Ottawa Co.: Stevens 2351 (an, on sheet with Stevens 2530, P. pubescens); 
Texas: Brewster Co.: Cory 35570, five and three-quarter miles east of Alpine, 
Sept. 19, 1940 (GH) is somewhat doubtfully referred to this taxon. 

20. Physalis Greenei Vasey and Rose, Contr. U. S. Natl. Herb. 1: 18. 
1890; P. pedunculata Greene, Pittonia 1: 268-269. 1899, non Mart. and 
Gal., Bull. Acad. Brux. 12: 132. 1842. 

Annuals, villous or short pilose, glandular and viscid; principal leaf blades 
2-4 em. long, ovate; petioles one-half as long as to slightly longer than the 
blades; leaf margins dentate to sinuate-dentate, or rarely entire; corolla 8-10 
mm. long, yellowish, or sometimes with a slightly darker tinge; anthers 1.5-2.5 
mm. long, yellow, on slender filaments; flowering calyx 3-4 mm. long on pedun- 
cles 15-30 mm. long; fruiting calyx 2-2.5 em. long, pointed-ovoid, on peduncles 
15-40 em. long. 

The smaller anthers of this species will serve to distinguish it, and 
separate it from P. crassifolia in those instances in which they tend to 
resemble each other. 

TYPE: Charles F. Pond Feb. 1889, Cedros Island, off the coast of Lower 
California; “southwest side of the island" according to Greene; type and 
isotype (us). 

P. Greenei grows on hills and sea-cliffs, southern California; it flowers in 
February, March and April. 


1958] Waterfall,— Genus Physalis in N. America 171 


Collections examined: CALIFORNIA: Orange Co.: Abrams June 12, 1901 
(ny); Mason 2933 (au, uc); Placer Co.: Jones 88 (GH); San Diego Co.: Abrams 
3309 (GH, NY, PH, UC, US); Allen 77 (GH); Jones March 1882 (au, uc); Wiggins 
1821 (uc). 


21. Physalis lobata Torrey, Ann. Lyc. Nat. Hist. New York 2: 226- 
227. 1828; Quincula lobata (Torr.) Rafinesque, Atlantic Journal, 1: 145. 
1832; P. sabeana Buckley, Proc. Acad. Sci. Phil. 14:6. 1863; Chamaesara- 
cha physaloides Greene, Bull. Torr. Bot. Club 9: 122. 1882; Quincula 
lepidota Aven Nelson, Bot. Gaz. 47: 430. 1909. 

Chamaesaracha physaloides is included here on the basis of Greene's 
phrase ‘‘flat seale-like hairs," which seems to be a good description of the 
appearance of the characteristic crystalline vesicles of P. lobata after they 
are dried. These structures are not found on P. Wrightii, the other species 
to which this name has been referred. 

Perennial, branching from the base, the branches spreading or procumbert; 
indument consisting of a varying amount of crystalline vesicles, flattening 
when dried, which may be abundant enough to give the plant a scurfy ap- 
pearance, or may be very sparse; principal leaves usually 4-10 em. long, with 
blades usually 0.5-3 em. wide, ovate-lanceolate to linear-lanceolate, cuneate 
at the base to a winged petiole, usually pinnatifid, rarely sinuate-toothed or 
entire; corollas blue or violet (rarely white), rotate, 1.5-2 em. broad, with five 
hairy pads on its base near the point of attachment of the filaments and alter- 
nating with them; anthers about 1.5-2 mm. long, yellow, on slender filaments; 
style twisted and bent to one side; flowering calyx 3-4 mm. long, its lobes 1.5-2 
mm. long, deltoid; flowering peduncle 1-3 (5) em. long; fruiting calyx 1.5-2 
em. long, pentagonal-ovoid, inflated, on peduncles 1-2.5 (3) em. long; seeds 
usually somewhat crenate on their backs. 


TYPE: James “On the Canadian"; not seen. 

P. lobata grows on plains, prairies, mesas, canyons, juniper barrens, 
desert areas and various disturbed habitats principally in western Kansas, 
Oklahoma and Texas, and eastern Colorado and New Mexico, and southern 
Arizona. 

Three hundred forty seven sheets of 275 collections have been seen from: 
Arizona, California (Fremont's Expedition in 1845), Colorado, Kansas, 
Nevada (Clark Co.), New Mexico, Oklahoma and Texas. 

A white-flowered forma is rarely found. It may be deseribed as P. lobata 
Torr., forma albiflora Waterfall, f. nov., corollis albis. TYPE: J.J. Thorn- 
ber Aug. 11, 1901, Experiment Station Range Reserve, Pima County, 
Arizona (Artz). Another collection is B. C. Tharp and C. Havard 49344, 
5-6 miles west of Del Rio, Valverde Co., Texas, April 16, 1949 (TEX). 

22. Physalis Carpenteri Riddell ex Rydberg, Mem. Torr. Bot. Club 4: 
330-331. 1896; P. Carpenteri Riddell, N. Orl. Med. and Surg. Journ. 
759. 1852, as a nomen nudum; Bot. Gaz. 3: 11. 1847 in synonymy. 


Plant over two-thirds of a meter tall, widely branched; herbage short-hairy, 
the leaf blades sometimes nearly glabrous and the calyces sometimes with a 
few long hairs; principal leaf blades (3) 7-11 cm. long, ovate to lanceolate- 
ovate, acuminate; petioles one-third to two-thirds as long as the blades; leaf 
margins usually entire, sometimes repand; flowers from single to 3-6 in the 
axils of the leaves; the fascicled flowers apparently due to the presence of a 


172 Rhodora [Vor. €0 


telescoped axillary branch, sometimes 2-4 em. long and bearing reduced leaves 
(Curtiss 6901: uc, GH, NY) as well as flowers; corolla about 1 em. long, yellow; 
anthers about 2 mm. long, yellow, on slender filaments; flowering calyx 4-5 
mm. long on peduncles 5-8 mm. long; fruiting calyx about 1.5 em. in diameter, 
nearly spherical, only slightly inflated; fruit having 1 to several plump, rounded, 
corky, seed-like bodies (possibly a peculiar development of unfertilized ovules) 
in addition to the normal, more or less reniform, flattened seeds. 

The species may be annual, according to some collectors, or from a deep- 
seated “rootstock” according to others; all the specimens examined were 
branches only. 

The author prefers to leave this species in Physalis, regardless of its 
peculiar characteristics, until a more thorough study of related genera, or 
possible subgenera, can be made. 

Collections examined: ALABAMA: S. B. Buckley, April (xy); FLORIDA: 
Columbia Co.: Geo. V. Nash 2503, Aug. 29-31, 1895 (an, Micn, Ny); Erdman 
West, seeds from Fort White, raised by Margaret Young Menzel as her 508a 
(TEX); Escambia Co.: Curtiss, 1886, Pensacola (GH); Suwanee Co.: A. H. 
Curtiss 6901, annual 2 ft. high and widely branched, growing in a cultivated 
field near Wellborn, Sept. 14, 1901 (Gu, Ny, uc); County undetermined: M. A. 
Curtis, Florida?; Loutstana: East Feliciana Parish: Riddell, March 1878 (an, 
one fruiting calyx); Orleans Parish: Drummond New Orleans (au); Ingals in 
1835, New Orleans (Ny); West Feliciana Parish: R. S. Cocks, 8603, common in 
rich woods (NY). DEPT. OF BOTANY AND PLANT PATHOLOGY AND THE RESEARCH 
FOUNDATION, OKLAHOMA STATE UNIVERSITY, STILLWATER, OKLAHOMA, 


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Bowpicn, THomas Epwarp. 1825. Excursions in Maderia and Porto Santo. 
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Buckuey, S. B. 1863. Descriptions of New Plants from Texas. Proc. 
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Curtis, M. A. 1849. New and Rare Plants, Chiefly of the Carolinas. Am. 
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Dr CawNpoLLE, AUGUSTIN Pyramus. 1813. Catalogus Plantarum Mons- 
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Frernautp, M. L. 1947. Additions to and Subtractions from the Flora of 
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Gray, Asa. 1875. Synopsis of North American Species of Physalis. Proc. 
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GREENE, E. L. 1882. New Western Plants. Bull. Torr. Bot. Club 9: 122. 

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HORNEMANN, JENS WILKEN. 1819. Hortus Botanicus Hafniensis, Supple- 
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Jacquin, Josera Franz. 1844. Ecologae Plantarum Rariorum. Vindo- 
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€ 


1958] Waterfall, —Genus Physalis in N. America 173 


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Lamarck, JEAN Baptiste. 1786. Encyclopédie Méthodique Botanique. 
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MACKENZIE, K. K. ann B. F. Buss. 1902. New Plants from Missouri. 
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Masters, MAXWELL TyrLpEN. 1894. Physalis Francheti. Gard. Chron. 
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Menicus, FRIEDRICH Casimir. 1780. Observationes Botanicae. Academia 
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MENZEL, MARGARET Y. 1951. The Cytotaxonomy and Geneties of Physalis. 
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MILLER, Parr. 1768. The Gardener's Dictionary. London: privately 
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NELSON, AvEN. 1909. Plantae Gooddingianae. Bot. Gaz. 477 425-436. 

NUTTALL, THoMas. 1834. Indigeneous Plants of the United States. Journ. 
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RYDBERG, Per AxEL. 1895. New Species of Physalis. Bull. Torr. Bot. 
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Torrey, Joun. 1828. Plants Collected During a Journey to the Rocky 
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WATERFALL, U. T. 1950. Some Results of a Third 5Summer's Botanizing in 
Oklahoma. Ruopora 52: 171. 


174 Rhodora [Vor. 60 


X Carex DkAMIL IN. Missovnr.— This interesting sedge was 
described by Hermann (Ruopora 40: 81. 1938) as a hybrid between 
C. Shortiana and C. typhina from Pike County, Indiana. The 
hybrid has thicker spikes (7-8 mm. wide) and longer beaks (1-1.5 
mm. long) of the perigynia than are found in C. Shortiana. 

It may now be recorded from Missouri on the basis of the fol- 
lowing three collections, all in the herbarium of the Chicago Na- 
tural History Museum: Allenton, St. Louis Co., July 30, 1887, 
G. W. Letterman (specimen on right hand side of sheet); swaley 
margin of shallow sinkhole pond on wooded upland, T 23 N, R 8 W, 
west part of sect. 15, 4 mi. south of West Plains, Howell Co., June 
25, 1955, Steyermark 78724; low wet woods in valley of Old Chariton 
River and bordering New Chariton River where swamp existed 
but is obliterated, T 62 N, R 16 W, SW l4 sect. 27, 214 mi. south 
of Youngstown, Adair Co., Sept. 19, 1955, Sleyermark 79705. At 
the last locality both C. squarrosa and C. typhina were present, but 
at the Howell County locality only one of the putative parents, 
C. Shortiana, assigned by Hermann, was present, represented by 
Steyermark 78725. Instead of C. typhina, the other putative parent 
assigned by Hermann, there was present C. squarrosa, represented 
by Steyermark 78723. It is interesting, therefore, to record a dif- 
ferent putative parent at the Missouri locality, i.e., C. squarrosa, 
rather than the one found by Hermann at the Indiana station for 
X C. Deamii. 

As this hybrid eventually becomes collected elsewhere, it will be 
interesting to learn which of the putative parents predominate. 
The morphological distinctions effected by the hybridization of C. 
typhina and C. Shortiana apparently cannot be differentiated from 
those effected by the union of C. squarrosa and C. Shortiana. While 
admittedly C. squarrosa and C. typhina are related species, sepa- 
rated chiefly by the spreading-divaricate vs. ascending beaks of 
the perigynia, one would expect some marked differences between 
the hybrids resulting from crosses of each one of these species with 
C. ShortianaJuLian A. STEYERMARK, CHICAGO NATURAL HIS- 
TORY MUSEUM AND MISSOURI BOTANICAL GARDEN, 


AN ALBINO Form or Dipsacus svrvEsTRIS.—While botanizing 
an undeveloped section of Mt. Hope Cemetery in Chicago, Mr. 
Karl E. Bartel discovered a colony of over fifty plants of a white- 
flowered Dipsacus sylvestris. Over one hundred heads of flowers 


1958] Gleason,— Two New Stations for Carex picta 175 


were counted. Only one stunted lavender-flowered plant was noted 
in the group. 

Since most other white-flowered plants are recognized with a 
formal name, it is consistent to provide a name for the present 
white-flowered teasel. 

Dipsacus sylvestris Huds., f. albidus Steyerm., forma nova. A 
forma sylvestris recedit corollis albidis.—Illinois: Mt. Hope Ceme- 
tery, 115th St., Chicago, Cook Co., Aug. 25, 1957, Karl E. Bartel 1, 
HOLOTYPZE, in Herb. Chi. Nat. Hist. Mus.—JULIAN A. STEYERMARK, 
CHICAGO NATURAL HISTORY MUSEUM AND MISSOURI BOTANICAL 
GARDEN 


Two New STATIONS FOR CAREX Picta. This rare sedge of 
anomalous structure was discovered more than a century ago 
near New Orleans by Drummond. Since then it has been de- 
tected in Winston County, Alabama, and in five adjacent counties 
in south-central Alabama. These are the only stations reported 
by Mackenzie in North American Flora (1935): if other stations 
in the southern states have been discovered in the last twenty 
years they are at present unknown to me. Few sedges have 
such a disjunct distribution, and these three widely separate 
areas suggest the possibility of its occurrence in suitable localities 
in Kentueky, Tennessee, and Mississippi. 

Carex picta steud. was collected March 18, 1955, by a stream 
in woods near Meridian, Lauderdale County, Mississippi, by 
G. R. Cooley, A. 5. Pease, and James D. Ray, Jr., number 3178. 
On March 29, 1956, Cooley and Ray collected it again in a wooded 
ravine opening into Tanyard Branch, north of Cross Road, 
Tishomingo County, Mississippi. These two stations are about 
175 miles apart, while the second one is about 75 miles northwest 
of the known Alabama station. Specimens will be deposited at 
the Gray Herbarium, the New York Botanical Garden, and 
Mississippi State College. 

The plant blooms early and must be very conspicuous at 
that time, due to its large clavate spikes with deep red scales. 
It is one of the few sedges which can be identified without 
perigynia. The plants are strictly dioecious and each flowering 
culm bears a single spike. The basal scale of the spike is some- 
what elongate and almost completely surrounds the rachis.— 
H. A. GLEASON, GREENWICH, CONN. 


176 Rhodora [Vor. 60 


A New BnvopruvrE FLora'. With this publication, A. LeRoy Andrews 
has filled a long existing lacuna in both bryology and plant. geography. 
The work is to bryologists, what the FLORA OF THE CAYUGA LAKE BASIN, 
NEW york, by K. M. WikGAND AND A. J. Eames is to students of the 
vascular plants. 

Begun by the author in 1908, the publication under review represents 
a thorough survey of a relatively small, but botanically important area. 
Dr. Andrews has collected most of the species found in the region by such 
botanists as Atkinson, Brewer, Brown, Cipperly, Dudley, Durand, 
Graham, Jackson, Kellerman, Nanz, Pratt, Rowlee, Schuster, Whetzel, 
Wiegand, and Winne. Several species not previously reported from the 
area have been found by the author and are treated in this flora. Only 
specimens actually seen by Dr. Andrews are definitely cited; any others 
are listed as “reported.” 

Since Wiegand and Eames, in their previously mentioned work, de- 
scribed the geological and physiographical features of the region in 
considerable detail, these matters are given brief treatment. 

The distribution and local site of each species is given as well as the 
names of collectors and dates of collection. 

In the matter of nomenclature, Dr. Andrews begins with Linnaeus’ 
SPECIES PLANTARUM of 1753 and observes the rule of priority “with some 
concessions in doubtful instances and no unfamiliar names." 

The keys, which should be of considerable help to students of New 
England bryophytes, are of excellent quality—brief and concise. A list 
of references pertaining to the bryophytes of the region is given. This 
work, a valuable piece of research, should be in the library of all readers 
of Ruopona, bryologists, and students of plant geography.—FRANK J. 
HILFERTY, DEPARTMENT OF BIOLOGICAL SCIENCES, MASSACHUSETTS STATE 
TEACHERS COLLEGE, BRIDGEWATER, MASS. 


! THE BRYOPHYTE FLORA OF THE UPPER CAYUGA LAKE BASIN, NEW YORK by A. LeRoy Andrews, 
Professor Emeritus of German and Honorary Curator of the Bryological Collections of the 
Wiegand Herbarium, at Cornell University. Cornell University Experiment Station, Cornell 
University, Ithaca, New York, 1957; 87 pp. 


Volume 60, No. 713, including pages 117-144, was issued 19 June, 1958. 


Clu qU ¢ f 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS l 
RALPH CARLETON BEAN 
CARROLL EMORY WOOD, JR, 


| Associate Editors 
IVAN MACKENZIE LAMB 


Vol. 60 July, 1958 No. 715 
CONTENTS: 


Perennial Ragweeds (Ambrosia) in Michigan, with the Descrip- 
tion of a New Intermediate Taxon. W. H. Wagner, Jr. and 
FB Soe T Tr A cet 177 


An Unusual Botanical Area in Missouri. Julian A. Steyermark 205 


The New England Botanical Club, Inc. 


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CARD-INDEX OF NEW GENERA, SPECIES AND 
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Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 July, 1958 No. 715 


PERENNIAL RAGWEEDS (AMBROSIA) IN MICHIGAN, 
WITH THE DESCRIPTION OF A NEW, 
INTERMEDIATE TAXON 


W. H. WAGNER, JR. AND T. F. BEALS! 


Michigan is a key state for the study of the eastern spread of 
Ambrosia coronopifolia Torrey & Gray, the so-called “western 
ragweed.” Present evidence indicates that what is generally 
‘alled the “western ragweed,” and usually identified as A. 
psilostachya DeCandolle in the literature of allergy and in many 
botanieal manuals, comprises actually a series of closely related 
but more or less distinguishable types which, for the purposes of 
this report, will tentatively be treated as species, as was done by 
Rydberg in 1922. These taxa differ from each other in charac- 
ters of hairiness, plant habit, distribution, shape of fruit, pollen 
size, and other characters, although the differences may be in 
statistical trends. The true or entirely typical A. psilostachya 
apparently does not grow in the central United States. The 
easternmost outlier of the complex, which is A. coronopifolia, is 
the sole species of ragweed in the central states known to be 
perennial, ie., to have underground vegetative reproduction. 
The investigation to be reported here grew mostly out of our 
curiosity as to what importance the perennial ragweeds have in 
forming the ragweed populations in Michigan. We have 
succeeded in adding a large number of new records to the known 
range of perennial ragweeds in this state and have compiled 
considerable new information concerning them. 

' Publication No. 5 on Atmospheric Pollution by Aeroallergens under research grant No. 


12-1379 (C) from the National Institute of Allergy and Infectious Diseases, Public Health 
Service. 


178 Rhodora (Vou. 60 


Ambrosia coronopifolia is the least abundant generally of the 
three ragweeds (A. artemisiifolia, A. coronopifolia, and A. trifida) 
heretofore known in Michigan (Lovell, Mathews, and Sheldon, 
1953). It has been reported to be “rare or absent" in the Upper 
Peninsula of Michigan, and “found occasionally to frequently” 
in the Lower Peninsula (Buchholtz et al., 1954). There is a 
question whether the species is native to Michigan at all, and 
this problem will be considered in some detail below. One of 
the peculiarities of perennial ragweeds which has been reported 
earlier is that the period of pollen production differs from that 
of A. artemisiifolia (syn. A. elatior), the common" or “low 
ragweed,” and we attempted to determine whether this was true 
of the populations in Michigan. We also endeavored to de- 
termine in the field whether there were any circumstances 
indicating differences in ecological responses between the two 
species. 

Quite early in the study it became apparent that the natural 
rariation of Ambrosia coronopifolia is fairly extensive; but as 
more and more populations were examined it turned out that 
not all the perennial ragweeds in Michigan belong to this species. 
On the contrary, a perennial species was found to be present 
that resembled both A. coronopifolia and the annual A. artemi- 
sitfolia. Such a plant, so far as we can determine, has not been 
described previously. The new ragweed has proved to occur 
over a large area, with stations in a number of counties; and it 
has also proved to be able to spread and form extensive popula- 
tions in a given locality. The discovery of this new form 
generated comparative field and laboratory studies on all three 
of these ragweeds, in terms of their vegetative characteristics, 
their chromosomes, pollen grains, and fruits. 

This investigation was carried out by the senior author in the 
years 1956 and 1957. He was joined by the junior author in 
the summer of 1957, and the latter added a large number of 
chromosome and spore studies. We are indebted for assistance 
to various people, especially J. M. Sheldon and E. W. Hewson, 
directors of the Project on Atmospherie Pollution, for stimulating 
this study to be made; Mrs. D. A. Beals, for collecting California 
ragweeds for comparison; H. H. Bartlett, for the latin diagnosis; 
R. W. Hanlin, for help on measurements; E. G. Voss, for aid on 


1958] Wagner and Peals,—Perennial Ragweeds (Ambrosia) 179 


historical matters; K. L. Jones, for his reading of the manuscript; 
and the curators of the following herbaria for lending specimens 
and looking up records: New York Botanical Garden, Michigan 
State University, University of Michigan, University of Minne- 
sota, University of California (Berkeley), U. 3. National Her- 
barium, and the Gray Herbarium. This report will deal with 
Ambrosia coronopifolia first, and will follow with a discussion of 
the new, intermediate species. 

OCCURRENCE: Ambrosia coronopifolia is capable of forming very 
large populations locally. Because the reproduction takes place 
largely by vegetative means’, a given area may be populated by 
the derivatives of only one or a few original mother plants. 
“Pure cultures" of a given variant may be distributed over 
hundreds of square yards, but they are clonal and represent 
offshoots of a single plant. Usually the spatial distribution of a 
given clone can be readily recognized because of subtle dif- 
ferences of color, hairiness, size, and shape between the indi- 
viduals of different clones. Two or more different clones may 
intermingle more or less intimately but their members remain 
completely distinguishable. In Mecosta County, for example, 
numerous plants of a fruiting form with a bright-green color 
were found completely intermixed with a nearly sterile form with 
a dull-green color. 

The environments in which perennial ragweeds thrive in 
Michigan are apparently always disturbed sites, the disturbance 
mostly of artificial nature. The annual species, A. artem?stifolia, 
also grows in such habitats; and where the perennial species 
occurs, the annual one is almost always near by. The latter is 
usually unable, however, to invade grassy fields. The annual 
ragweed is generally confined to open soil, but the perennial 
species will readily invade adjacent grassy fields and grow 
competitively with fairly dense field vegetation. In this con- 
nection, one of the most interesting occurrences of A. corono- 
pifolia and A. artemisiifolia growing together was observed on 
the outskirts of Gaylord, Otsego County; here both species were 
scattered abundantly along the roadsides, but A. coronopifolia 


‘ 


2 The underground axes of the perennial ragweeds are commonly referred to as “rootstocks” 
or “rhizomes,” but this does not seem to be accurate. The reproductive organs appear to be 
roots anatomically rather than stems (as was correctly given by Fernald, 1950), and it is hoped 


that a detailed report on the process of reproduction will be published in the future. 


180 Rhodora [Vor. €0 


had spread over some sixty square yards of a large, mowed, 
private lawn, and its pale-green foliage was very conspicuous 
even though its shoots had been cut to the level of the grass. 
Only a few, scattered plants of A. artemisiifolia were seen in this 
lawn. 

In behavior A. coronopifolia is decidedly “weedy” in Michigan, 
as noted by the earliest writers who mentioned the species in the 
state. Of the state’s collections on which environmental data 
are recorded, about one-sixth specify that the habitat was along 
railroads, and one-third say “along roads" or “along highways." 
Over two-thirds of the collections came from about cities, villages, 
and settlements. The species can become a locally serious weed 
in yards and gardens (e.g., around the Interlochen R. R. Station 
area, and on the outskirts of Frederic and Gaylord). Because 
of its tendency to form large and dense clones it could provide a 
local hayfever problem, especially in the weeks before the annual 
species comes into flower, as will be discussed below. 

The soil in which A. coronopifolia grows is ordinarily dry, 
very well drained, and commonly sandy or gravelly. Nearly 
one-third of the labels on Michigan herbarium specimens use the 
words 'sandy" or “gravelly.” The habitats that are found 
away from roadsides and railway lines are generally clearings, 
“deserts,” open pastures, mowed fields, grass-covered hills, 
baseball diamonds, and waste ground and dumps. The species 
evidently can not withstand any great degree of shading; when 
in wooded areas it will always be discovered on the cleared ground 
along open trails, or in more or less exposed, prairie-like spots 
such as the open sandy and grassy hillsides in jackpine (Pinus 
banksiana) regions. 


Although A. coronopifolia may be extremely abundant locally, 
the species must be rated as only frequent to uncommon taking 
the state in its entirety. In the northern part of the state, i.e., 
the northern half of the lower peninsula and all of the upper 
peninsula, it may be found readily by driving along roads and 
highways, but this impression is misleading with respect to the 
state flora as a whole, because it is exactly those places where 
one is likely to drive where the species is most surely to be found. 
Ambrosia artemisiifolia is infinitely more common in the state as 
a whole. The giant ragweed, A. trifida, is locally frequent to 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 181 


common in the southern half of Michigan, but it becomes very 
rare in most of northern Michigan except around certain towns. 
Thus, where A. coronopifolia is the most common, A. trifida is 
usually rare or absent. In our field surveys, all three species of 
ragweeds were found growing together only at Cheboygan, 
Cheboygan Co., and Marquette, Marquette Co., in weedy city 
lots. Ambrosia trifida was considerably less common at these 
two places than the other ragweeds. 

There seems to be a question whether A. coronopifolia was 
originally indigenous in Michigan or not. Fernald (1950) gave 
the range of this plant (as A. psilostachya var. coronopifolia) as 
“Mich. to Sask. and Mont., s. to La., Tex., and Mex.; adv. e. to 
Quebec, N.S. and N.E.” On the contrary, Cronquist in Gleason 
(1952) circumscribed the range (of A. psilostachya) as “Ill. to 
La., w. to Sask., Ida., Cal., and n. Mexico; introduced eastward," 
and thus left Michigan out of its presumed original range. 

Today A. coronopifolia is widespread in Michigan and it has 
been recorded from forty-three counties, as shown in Figure 1, 
A. The species extends across the Upper Peninsula down to the 
middle of the Lower Peninsula. Further southward, i.e., in the 
bottom half of the Lower Peninsula, the species is frequent only 
in the western or Lake Michigan side of the state. It is ex- 
ceedingly rare in the southeastern quarter of the Lower Peninsula 
at the present time, and we have not succeeded in finding any 
populations; there is only one record from this part of Michigan. 

The earliest definite record of this species in Michigan known 
to us was in the year 1900, and there are apparently no prior 
collections’. The first mention of perennial ragweeds in the 
state was in a list published by Daniels (1904) of plants found at 
Manistee, Manistee Co., that had not been included in Michigan 


’ That it might have been collected as early as September, 1831, by Douglass Houghton, 
was suspected because of two herbarium sheets so dated and labelled “Fox River of L. 
Michigan" (MICH). The notion was dispelled, however, by consulting H. R. Schoolcraft’s 
description (1834) of his expedition in 1831. Schoolcraft wrote that “At Galena [Illinois] the 
exploring party separated, part returning in canoes up the Wisconsin, and part crossing the 
mine country, over the branches of the Paktolika, and by way of the Blue Mounds, to Fort 
Winnebago. From this point, Fox River was descended to Green Bay, and the route of the 
lake coast pursued northward to the straits, and to the Sault of St. Mary." Thus the "Fox 
River" of Houghton's 1831 collections of A. coronopifolia came from the river of that name in 
Wisconsin, south of Green Bay. It is interesting that Houghton himself (in Schoolcraft, op. 
cit.) gave the locality for various plant species of his report as “Fox River, N.W. Terr.," but 
did not list this species. 


182 Rhodora [Vor. 60 


by previous writers. He lists the species under “Weeds” as 
occurring in “Yard and roadside, Maple Street, near Catholic 
Cemetery," and a specimen from this collection is deposited in 
the Michigan State University Herbarium. For the two annual 
species of ragweeds, A. artemisiifolia and A. trifida, there are 
much earlier records: A. trifida was taken as early as 1838 by 
Houghton’s survey at White Pigeon, St. Joseph Co. (and there 
are other collections as early as 1861 and 1869). Ambrosia 
artemistifolia was obtained in the same year, by the same survey, 
but in Cass County (MICH). 

That A. coronopifolia could have been overlooked during the 
entire nineteenth century seems quite unlikely unless the species 
was exceedingly rare. Its usually large clones with their charac- 
teristic pale foliage are readily noticed in the field. Ambrosia 
coronopifolia was collected considerably earlier in the states to 
the west (namely Wisconsin, Illinois, South Dakota, and Minne- 
sota) as shown by specimens in the herbaria of the University of 
Michigan, Michigan State University, and other institutions. 
Many parts of Michigan, where A. coronopifolia now forms a 
conspicuous element of the vegetation were reasonably well 
collected by botanists prior to 1900. The region of Douglas 
Lake (Emmet and Cheboygan Cos.), for example, was examined 
by a number of botanists during the nineteenth century (Voss, 
1956), but none of them found the perennial ragweed. 

In 1899, C. K. Dodge completed a flora of St. Clair Co. based 
on nearly twenty-five years of collecting, but in a list of 1,112 
species A. coronopifolia was lacking. | Four years later he dis- 
covered it for the first time in this county. Dodge, who was a 
very active observer and collector of Michigan flora, and con- 
tinued his work through the early decades of the present century, 
evidently concluded that this species was in the process of 
becoming naturalized. His statements, such as “a weed noted 
in waste places of Marquette, Negaunee, and Ishpeming... 
becoming frequent" (1918); and “becoming established in cities 
and villages as a perennial weed" (1921) clearly indicate that he 
considered the species to be behaving as an adventive. 

Professor H. H. Bartlett, another botanist of long experience 
with the Michigan flora, expressed his opinion recently (1952) as 
follows: “This is a prairie species that seems to be taking ad- 


1958] Wagner and Beals,— Perennial Ragweeds (Ambrosia) 183 


Fig. 1. County distribution maps of perennial ragweeds in Michigan: A. Ambrosia coro- 
nopifolia. B. A. X intergradiens. (Base maps courtesy of Cranbrook Institute of Science). 


vantage of the clearing of forest and making headway in an 
eastern invasion fairly well to the north but not in southern 
Michigan." Observations by him and Dr. C. D. Richards (both 
botanists allergie to ragweed pollen) on a field trip to the north 
in 1951 indicated that “north of Gratiot County, Michigan, the 
amount of ordinary ragweed decreased appreciably, with re- 
sultant relief from allergic symptoms." When they reached 
Emmet Co., they observed that “although we had run beyond 
the region of greatest abundance of the ubiquitous common 
species, Ambrosia arlemisiifolia, we had come into the newly 
extending range of another, namely A. psilostachya [Le., A. 
coronopifolia]." 

All that we can surmise, therefore, is that A. coronopifolia has 
spread since 1900 into a great number of localities in Michigan. 
It was either very rare and local, or non-existent, in the state 
prior to that time. It seems not unlikely that the bulk, if not 
all, of the present-day populations may have been introduced 
from further west—Minnesota, Wisconsin, and perhaps Illinois— 
where the species was in all likelihood native and well established. 
Deam (1940) considered this plant to be a rarity in nearby 
Indiana and to be introduced there. Moss (1956) has concluded 
on grounds similar to ours that another species, A. arlemisiifolia, 
had been introduced into southeastern Alberta and adjacent 


184 Rhodora [Vor. 60 


Saskatchewan, and he cites the fact that the earliest collection 
was in Saskatchewan in 1879. 

Surely the development of great railroad lines and innumerable 
roads and villages and towns over the state of Michigan during 
the last century has opened the way for active invasion of A, 
coronopifolia into areas where it never before existed. In the 
course of its spread it has come repeatedly into contact with the 
abundant and weedy A. artemisiifolia which has itself also 
inereased in numbers, and, where the two species have crossed, 
new types of perennial ragweeds have been generated which 
combine the characteristics of both, to be discussed below. An 
enumeration of present county distribution is given in the 
following list which contains the earliest records for 4. coro- 
nopifolia, the years indicated in parentheses: 


Alcona (1957), 1 mi. E. of Mikado, Wagner 8482 (uicu); Alger (1957), 
Melstrand, Wagner 8438 (mica); Allegan (1950), Ely Lake, Bazuin 8313 
(msc) also referred to by Kenoyer, 1934; Alpena (1957), Alpena, Wagner 
8483 (micu); Antrim (1956), W. of Alba, Wagner 8340 (Mic); Baraga 
(1950), N. of Baraga, Richards 4342 (micu); Barry (ca. 1930), reported 
by Bazuin, ms.; Benzie (1956), W. side of Co. 669, E. edge of Sect. 25, 
Wagner 8334 (mica); Charlevoix (1957), town of Walloon Lake, Wagner 
8434 (micn); Cheboygan (1913), Indian Settlement, Barnum (umes); 
Chippewa (1935), Sugar Island, Hermann 7235 (msc, Ny, vs); Clare 
(1957), Meredith, R.3W, T.20N, Sect. 13, Wagner 8513 (micu); Crawford 
(1956), S. border of Frederic, Wagner 8346 (micu); Delta (1949), 2 mi. E. 
of Rapid River, McVaugh 11170 (mich, CRANBROOK); Dickinson (1951), 
2.4 mi. W. of Norway, Bartlett & Richards 793 (micu); Emmet (1921), 
W. of Pellston, Ehlers 1817 (micu, umes); Gogebic (1919), 3-4 mi. N.E. 
Watersmeet, B. & D. 2779 (micu); Grand Traverse (1956), Interlochen, 
Wagner 8336 (micu); Houghton (1926), Calumet Water Works, Wolff 
795 (mic); Kalamazoo (1937), 6 mi. W. of Schoolcraft, Hanes 3827 
(NY); Kalkaska (1956), M-72, just E. of Co. 597, Wagner 8337 (micu); 
Kent (ca. 1930), reported by Bazuin, ms.; Keweenaw (1910), W. S. 
Cooper 274 (Gu); Lake (1948), 7 mi. W. of Baldwin, McVaugh 9797 
(mic); Leelanau (1956), E. of Empire, Wagner 8332 (micu); Mackinac 
(1913), Bois Blane I., C. K. Dodge (micu); Manistee (1900), F. P. Daniels 
s. n. (Msc); Marquette (1916-17), reported by Dodge, 1918, collected 
along M-28 at Marquette, Wagner 8440, in 1957 (mica); Mecosta (1957), 
0.3 mi. N. of Stanwood, Wagner 8499 (mich); Menominee (1933), Grassl 
2623 (Mich); Missaukee (1957), Lake City, Wagner 8475 (micu); Montcalm 
(1957), 2 mi. N. of junct. U.S. 131 and M-46, Wagner 8506 (mcn); 
Muskegon (1949), Cedar Creek Twp., Sect. 18, Bourdo 25 (micu); Newaygo 
(1916), Bessey & Darlington 1106 (msc); Ogemaw (1957), West Branch, 
Wagner 8480 (icu); Osceola (1957), 3 mi. N.W. junction of M-61 and 
M-115, Wagner 8500 (icu); Otsego (1956), S. of Gaylord, Wagner 8344 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 185 


(MicH); Presque Isle (1949), Bearinger Twp., Marshall 857 (msc); 


H 


Roscommon (1919), N. end of Higgins Lake, Bessey s.n. (Msc); St. Clair 
(1903), Port Huron, C. K. Dodge (micu); Schoolcraft (1915), reported by 
Dodge, 1921; Van Buren (1906), very rare, Pepoon 944 (msc); Wexford 
(1956), juncture M-115 and U.S. 131, Wagner 8330 (mica). 


PERIODICITY: Wodehouse (1945, table iv) has already indicated 
that the pollen of “Ambrosia psilostachya” precedes that of A. 
artemistifolia in the atmosphere by two weeks. It was therefore 
no surprise that field studies of the two species in Michigan 
revealed the same approximate relationship between A. coro- 
nopifolia and A. artemisiifolia in their morphological develop- 
ment. A total of 338 plants were collected during the period 
August 3—5, 1956, 234 of A. coronopifolia from eight localities in 
eight counties, and 104 of A. artemisiifolia from nine localities in 
nine counties. These specimens were then measured for length 
of the staminate spike primordia. In A. coronopifolia, 18% of 
the total branches bore staminate spikes which were over 3 em. 
in length, many of these with mature flowers. In A. artemi- 
sitfolia, a mere 1% of the total bore spikes of such development 
during the first week of August. In A. coronopifolia, only 19% 
of the total number of branches lacked visible spike primordia, 
but in A. artemisiifolia 64% had no visible primordia. About 
one week earlier the same year, Professor K. L. Jones reported 
that on July 27-28, he developed a hay-fever reaction while 
visiting the city of Cheboygan. Along the estuary there, he 
discovered a large number of plants of A. coronopifolia in anthe- 
sis, while the A. artemisiifolia was not yet in bloom. The 
earliest herbarium specimen with flowers in A. coronopifolia in 
Michigan is July 12 (Menominee, C. O. Grassl 2623, 1933, 
MICH). 

VARIATION: Of the three basie ragweed species in Michigan, 
A. coronopifolia seems to be the least variable. The uniformity 
of any given stand, however, may be misleading, since the indi- 
viduals are clonal and tend to be derived from one or a few 
original plants as discussed earlier. The most conspicuous 
variations include those in leaf form, leaf arrangement, plant 
habit, and fruit characteristics. 

The leaves have been described by Rydberg (1922) as “ovate 
in outline, pinnatifid, subsessile, or the lower with short winged 
petioles.” The outline may vary, however, as shown in Figure 2 


186 Rhodora [Vor. 60 


4 


Fig. 2. Outlines of approximately median leaves of moderate-sized plants of ragweeds from 
Michigan (all petiole bases not complete): A. Ambrosia artemisiifolia: 1. Washtenaw Co.; 2 
Otsego Co.; 3. Benzie Co.; 4. Crawford Co. B. A. X tntergradiens: 1. Crawford Co.; 2. Benzie 
Co. C. A. coronopifolia: 1. Benzie Co.; 2. Grand Traverse Co.; 3. Leelanau Co.; 4. Antrim 
Co.; 5. Otsego Co.; 6. Kalkaska Co. 


(e.g., 2 vs. 3) from approximately lanceolate to deltoid. Ex- 
tremely large leaves (not figured) from plants growing under 
luxuriant conditions may approach in outline and cutting the 
much smaller, typical leaves of A. artemisiifolia growing under 
normal conditions. The length of the petiole diminishes from 
the basal to the median leaves, varying from obviously petiolate 
to sessile, making comparisons quite difficult from collection to 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 187 


collection unless leaf position is taken into consideration. The 
dissection of the leaves ranges from coarsely lobed to nearly 
bipinnatifid. In the least dissected extreme the leaf-blades are 
long-ovate and the sinuses so shallow that they extend only one- 
third to one-half of the way from the blade margin to the midrib. 
The other extreme is represented by triangular leaf-blades cut 
seven- to nine-tenths of the way to the rachis, resembling in leaf 
outline A. artemisiifolia forms except for the relative length of 
the petiole. The blade/petiole ratio of median leaves averages 
4.2 but varies from 3.1 to 8.0. 

Occasional specimens of A. artemisiifolia are collected, es- 
pecially in southern Michigan, which resemble A. coronopifolia 
in foliar characters. Generally, however, the leaves of the latter 
may be distinguished by the following ensemble of differences, 
some of them subtle and not readily evident on the herbarium 
sheet: (a) thicker leaf texture; (b) harsher, more appressed hairs; 
(c) paler green color; (d) fewer lobes and segments; (e) broader 
midrib wing; (f) shorter petiole; and (g) narrower blade outline. 
All these characters tend to overlap individually to some extent 
between the two taxa, and they must, therefore, be considered 
together in making identifications. 

Leaf arrangement in Ambrosia coronopifolia does not differ 
basically from that of the other ragweeds. Depending on size 
of plant, the lower four to nine leaf pairs are opposite, but the 
leaves above and especially those from the axils of which the 
staminate inflorescences arise tend to be subopposite to alternate. 
At Interlochen, Grand Traverse Co., however, there is at least 
one large clone, growing with clones of the ordinary opposite- 
leaved form, in which the leaves are all whorled, with three 
leaves per node. In this “leafy” form, the spacing of the nodes 
is like that in the typical form, and in the upper part of the 
plants the whorled condition gives way to the usual alternate- 
leaved state. 

In habit the plant is usually a simple leafy axis. The vast 
majority of plants in typical exposed situations on sterile soil 
along roadsides and railways have simple axes with only a 
single terminal staminate spike or with one or a few laterals in 
the upper fourth of the plant. A few more lateral branches may 
tend to develop from axillary buds as the season progresses, thus 


188 Rhodora (Vor. 60 


spreading the potential flowering time. If the main stem of a 
plant is eut off or otherwise damaged during the first half of the 
summer, a short, “bushy”? specimen will result, numerous 
branches arising from the base of the stem. 

'The only profusely branched forms that are not the result of 
injury are the giant plants found in unusually rich environments. 
This large form was found at the edge of a vacant lot in Alpena, 
Alpena Co. (Wagner 8483) and on a farm near the juncture of 
highways M-72 and Co. 597 in Kalkaska Co. (8337). At the 
latter site, the normal form with few branches occurred in fields 
and had spread into open soil in a cultivated truck garden where 
giant plants up to 70 em. tall formed, bearing in some cases over 
30 large and well developed staminate spikes with numerous 
coarse vegetative branches arising from near the base. "These 
were very likely stimulated to grow and branch by the unusually 
rich soil conditions in the truck garden where they were free of 
competition. In growth habit they resembled the large, branch- 
ing plants characteristic of true Ambrosia psilostachya as it grows 
in California, rather than the simpler forms typical of A. coro- 
nopifolia as it occurs in the Great Lakes area. 

'The morphology and maturation of fruits varies to some extent 
from clone to clone. It is not at all uncommon to find popula- 
tions of A. coronopifolia in which only a small number of fruits 
have been produced. Such populations may co-exist side by 
side with others which have fully developed fruiting. Morpho- 
logically the fruits of A. coronopifolia usually have very short 
lateral processes or none at all so that they become, in the 
extreme form, entirely rounded at the top except for the beak. 
One striking population .at Yuma, Wexford Co. (8497a), however, 
has fruits with rather conspicuous processes, suggesting the fruits 
of A. artemisiifolia. 

POLLEN GRAINS: The pollen grains of Ambrosia coronopifolia 
were studied in terms of size, and presence or absence of proto- 
plasts. To measure diameters, anthers were removed from 
herbarium specimens, placed in 75% aqueous ethyl aleohol on a 
microscope slide and glycerine jelly containing acid fuchsin was 
added. The measurements were made of the widest diameters 
of 20 grains for each collection. No broken or collapsed grains 
were measured. To estimate the number of inviable grains, the 
technique was to crush dried anthers in aceto-carmine and to 


1955| Wagner and Beals,—Perennial Ragweeds (Ambrosia) 189 


Fic. 3. Fruits of ragweeds collected near Stanwood, Mecosta Co., Mich., September 28, 1957 
(specimens drawn without trichomes): A. Ambrosia artemisiifolia (from several plants). 
B. A. X intergradiens (each horizontal row from a different clone). C. A. coronopifolia (each 
row from a different clone). 


heat the freed pollen grains until all the protoplasts became 
stained. "Those grains in which no protoplasts were evident by 
staining were counted and compared with those in which the 
protoplasts did stain. Approximately 1000 grains were recorded 


190 Rhodora [Vor. 60 


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ARR | 403 
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PP 2 AN fun? \ 
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CX | mt + | ( tr / 4e o ^ 
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$e ? OT d \ zT, Md / M 
(s TIN m Nee fo | d a 
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sue — Hu €. 0$ 
DON ^ (C y - 
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5 ~ B a 10 p / , ~ 
Fic. 4. Chromosomes of Michigan Ambrosia. 1-5. A. coronopifolia, 2n = 72: 1, Antrim 


Co., metaphase II, 8340; 2, Emmet Co., metaphase I, 8446; 3, Benzie Co., metaphase 1, 8334; 
4, Mackinac Co., metaphase I, 8435; 5, Alger Co., mitotic metaphase, 8438-bl. 6-11. A. X 
intergradiens, 2n = 54: 6, 9, Emmet Co., metaphase II, 8445; 7, 8, Emmet Co., late prophase, 
8445; 10, Alger Co., mitotic metaphase, 8438-2; 11, Emmet Co., prophase I, 8445. 12. A. 
psilostachya, 2n = ca. 108, Los Angeles Co., California, metaphase II, D. A, Beals. 


in this way with mechanical counters, and the results expressed 
as percentage abortive grains of the total. 

The average pollen diameter of 11 collections of Ambrosia 
coronopifolia is 20.5 microns, with a range in individual grains of 
17.6 to 25.0, and a standard deviation of 1.05. An average of 
22.6% of the pollen grains from 11 collections appeared to lack 
protoplasts, with a range from 5 to 72%. Comparison of these 
values with those of the other ragweeds involved in this report is 
shown in Table 1, and will be discussed below. 

CHROMOSOMES: To determine chromosome numbers, specimens 
were fixed in Newcomer's fluid (Newcomer, 1953). For meiotic 
observations, whole young staminate spikes were placed in the 
fixative after removal from plants grown in the University of 
Michigan Botanical Gardens (August, 1956), and from the wild 
(July, 1957). At the time of examination, individual involucres 
were removed, the florets dissected out, and the anthers carefully 
removed and crushed in aceto-earmine stain on a microscope 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 191 


slide. The pollen mother cells were thus extruded, and were 
then squashed in the ordinary way. For somatic observations, 
root tips were removed from greenhouse plants and pre-treated 
before fixation in a saturated cool solution of paradichlorobenzene 
and kept in a 40? cold-room for 3-5 hours to shorten the chromo- 
somes. 'The roots were then fixed and squashed. Division 
figures were drawn using the camera lucida. 

As stated in a preliminary report based on observations made 
during 1956 by the senior author (Wagner, 1957), there are 36 
bivalents in meiotic metaphase, all of approximately the same 
size. This number has been confirmed by studies made in 1957 
by the junior author, and also by the report, based presumably 
on the same species, of Mulligan (1957). Meiosis in Ambrosia 
coronopifolia is evidently regular, and the division figures of all 
stages appear to be normal (Figure 4, 1-5). Counts of n = 36 
or approximately that number were made in meiosis of the 
following collections: Benzie Co., 8334; Antrim Co., 8340; 
Emmet Co., 8430; Mackinae Co., 8435; and 8444; and Marquette 
Co.; 8439. Counts of 2n — 72 or near that number were made 
in mitosis in the following collections: Kalkaska Co., 8337; 
Grand Traverse Co., 8336; Benzie Co., 8334; Alger Co., 8438bl; 
and Emmet Co., 8444-1. Representative specimens of all these 
collections are on deposit in the University of Michigan Her- 
barium. The collections from which the majority of meiotic 
numbers were determined were collected mainly on July 19, 20, 
and 21, 1957. 

INTERMEDIATE PERENNIAL RAGWEED: The second type of 
perennial ragweed in Michigan is easily overlooked by the 
collector because of its resemblances both to Ambrosia corono- 
pifolia and the annual A. artemisiifolia. In all respects, this 
plant is an intermediate between these two species, and it will 
therefore be described as a hybrid taxon, A. X zntergradiens'. 
The major characters of the new perennial ragweed are sum- 
marized in Table 1, in comparison with its relatives. In its 
habitat it is generally found with the two other ragweeds, but 

1 Ambrosia X intergradiens Wagner, hybr. nov. Perennis, multiplicatione vegetative colonias 
uniformes formans sed inter se multiformes. Ambrosiae coronopifoliae similis sed differt 
plerumque pilis numerosis valde divergentibus in caule primario, foliis plus dissectis, tenuioribus, 


petiolis longioribus, chromosomatibus (2n) 54, divisione meiotica irregulari. Communis in 
locis dispersis ruderalibus michiganensibus borealibus cum A. artemisiifolia et A. coronopifolia. 


192 


TABLE I. A COMPARISON OF THREE RAGWEEDS IN 


Duration 


Petiole length 
(em.) (ea. 30 
median lvs.) 

Number of pro- 
jections on bas- 
al segment pair 
(ca. 30 median 
lvs.) 

Narrowest rachis 
wing width be- 
tween two bas- 
al segment 
pairs. (ca. 30 
median lvs.) 

Indument of main 
stem and peti- 
oles (subjective 
judgment) 

No. pistillate fls. / 
cluster 

Per cent good 
fruits/cluster 

Terminal beak 
length of fruit 
(mm.) 

Length of 
“spines” (mm.) 

Per cent abortive 
pollen per col- 
lection 

Pollen grain di- 
ameter: over-all 
range (microns) 

Pollen grain di- 
ameter: aver- 
ages (microns) 

Pollen grain di- 
ameter: stand- 
ard deviation 

Chromosome 
number (27) 

Meiotic meta- 
phase 


| ay gy. 
| artemisiifolia 
| 


| 14.3-20.8 


Rhodora 


I 


Annual 


1.5 (0.3-3.0) 


9.0 (0-22) 


1.4 (1.0-2.0) 


Sparse and 
spreading (ex- 
cept f. villosa) 

5.4 (2-15) 

77 


1.2 (0.8-2.0) 


0.5 (0.2-0.8) 


16.0 (3-44) 
(9 colleetions) 


17.6 (16.3-19.7) 


0.79 (0.58-0.94) 


36 


Regular 


X intergradiens 


Perennial 


1.0 (0.5-1.7) 


5.8 (2-10) 


Mostly denser & 
spreading. 

3.1 (1-8) 

16 


0.9 (0.8-1.0) 


0.4 (0.2-0.6) 


54.9 (42-83) 
(12 collections) 


16.6-29.6 


21.8 (20.4-23.6) 


2,30 (1.36-3.22) 


54 


Irregular 


[Vor. 60 


MICHIGAN. 


coronopifolia 


Perennial 


0.9 (0.5-1.4) 


2.2 (0-10) 


3.1 (2.0-5.0) 


Denser and 
appressed. 


1.2 (1-3) 

55 

0.6 (0.5-0.8) 
0.2 (0.0-0.5) 
22.6 (5-72) 

(11 collections) 
17.6-25.0 

20.5 (19.9-22.2) 


1.05 (0.72-1.37) 


72 


Regular 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 193 


this is not always so. For collectors in general it is probably 
most profitable to compare the new plant with the other perennial 
species, A. coronopifolia. Ambrosia X intergradiens may be 
distinguished from the latter by its more spreading, more 
delicate hairs on the stem axis, these usually more abundant. 
The whole plant of the intermediate is slightly greener (not 
glaucous). Corresponding leaves tend to be more divided and 
they have narrower wings between the pairs of lobes. Corre- 
sponding leaves (i.e., those in like position on plants of approxi- 
mately equal size) will also tend to have slightly longer and 
narrower petioles. If the fruits have matured, the lateral 
processes and the beak will be more strongly developed than in 
A. coronopifolia so that the fruit is more suggestive of that in 
A. artemisiifolia. 

The hybrid perennial ragweed has turned out to be surprisingly 
numerous in Michigan. In 1956 and 1957 it is estimated that 
between forty and fifty different populations, large and small, 
have been discovered. "These were found in over twenty localities 
in fifteen counties. All collections in the following list were 
made by the senior author with the help of others, especially 
J. A. Churchill, R. F. Blasdell, and P. J. Neihaus. Unless 
otherwise indicated, both parental ragweeds were present at each 
station listed: 


Alger, weedy edges of roads at Munising Falls, Munising, 8438-2; Antrim, 
no parents within at least 100 yds., grassy plains along U.S. 131, 1.2 mi 
N. of county line, Sect. 31, R.6W, T.29N, 8491; Benzie, on steep road- 
banks, W. side of Co. 669, E. edge of Sect. 25, R.14W, T.26N, 8335; in 
overgrown vacant lot, Bendon, Sect. 23, T.26N, R.13W, 8493; Charlevoix, 
vacant lot in town of Walloon Lake, 8485; along R.R. tracks in Boyne 
City, 8487; Cheboygan, vacant lots, Cheboygan, 8434; Clare, Meredith, 
R.3W, T.20N, Sect. 13, 8513; Crawford, lawns, gardens, fencerows, S. 
border of Frederic, 8348; gardens and roadsides behind P.O., 8448; 
Emmet, sandy vacant lot by Greyhound Bus Station, Mackinaw City, 
8444-2; open grassy places along R.R. tracks, Pellston, $445; Grand 
Traverse, one or both parents present, in 3 places—(#1) in sandy lot near 
Elementary School, an all-pistillate clone, (72) just S. of bridge over 
Little Betsey Creek, and (73) by the R.R. Station, Interlochen, 8473; 
only A. artemisiifolia present, along railroad, 0.6 mi. W. of Grawn, Sect. 
12, R.12W, T.26N, 8492; Kalkaska, weedy fields at intersection of U.S. 
131, 1.5 mi. S. of Co. line, Sect. 2, R.7W, T.28N, 8488X; Marquette, both 
parents plus A. trifida present, vacant lot along M-28, Marquette, 8443a 


194 Rhodora [Vor. 60 


and 8443b; Mecosta, prairie-like fields, hybrids in sandy, more or less 
shaded, areas, 0.3 mi. N. of Stanwood on U.S. 131, 8498; Osceola, only 
A. artemisiifolia seen, grassy fields near R.R. Station, Marion, 8429; 
Otsego, 1.2 m. N.N.W. of Vanderbilt on M-27, 8515; Wexford, grassy 
fields at junction of M-115 and U.S. 131, 8474; along R.R. tracks, Yuma, 
Sect. 34, R.12W, T.23N, 8496a. 


The different intermediates are quite variable among them- 
selves, and it is conceivable that there is some degree of intro- 
gression involved in the formation of Ambrosia X intergradiens. 
In general, however, we have assumed that we are dealing with 
Fı hybrids which have become established and by means of their 
perennial habit and reproductive method have been able to 
form more or less large populations. It is interesting to note 
that A. X intergradiens is occasionally found with only one 
parent in the immediate neighborhood of the colony. At one 
place (Antrim Co.), in fact, we discovered a large population 
where careful searching failed to reveal either parent within at 
least 100 yards. The hybrid has been found to invade plowed 
fields (Otsego Co.), gardens (Crawford Co.), and plantations of 
pine trees (Benzie and Mecosta Cos.). The best localities for 
locating large populations are in grassy places along railroad 
tracks and sandy roads in and around towns and villages. 
Disturbed areas where both A. coronopifolia and A. artemisiifolia 
occur in a large mixture are almost sure to reveal at least one or 
a few populations of A. X tntergradiens. Further field studies of 
Michigan weeds will unquestionably yield many more populations 
of the new ragweed (once its characteristics are recognized), 
judging from the readiness with which we have found it to date. 

The major variations of A. X intergradiens include degree of 
hairiness and the size of the plant. The former is probably 
genetically controlled, and is perhaps determined more by 
inheritance from the annual ragweed, A. artemisiifolia, than the 
other parent. This is suggested by the fact that A. artemisiifolia 
is strongly variable in hairiness, and the form villosa Fern. & 
Grisc. is common throughout this area, contrasting by its dense, 
spreading-villous indument with the ordinary form. Ambrosia 
coronopifolia does not vary nearly so much in this respect. The 
size (i.e. , height and number of branches) of the plant, on the 
other hand, is probably controlled mainly by the environment: 


Rhodora Plate 1234 


WOH WAGNER 


PLATE 1234. Ambrosia X intergradiens W. H. Wagner, type specimen, 


1958] Wagner and Beals,— Perennial Ragweeds (Ambrosia) 195 


those growing in completely exposed sites tend to be smaller and 
to have a simple form with only one terminal, staminate raceme 
(Pellston, R. R. tracks, Emmet Co., 8445); those growing in 
more or less shady and damper sites tend to be larger and more 
“bushy” and to have numerous staminate racemes branching out 
below the terminal one (from near Stanwood, at edges of pine 
plantation, Mecosta Co., 8509-5). 

One of the most curious variants of Ambrosia X intergradiens 
was found at Interlochen, Grand Traverse Co. (8473-1). An 
entire clone was observed to comprise pistillate plants only. 
The pistillate flowers of these plants are borne in racemes in a 
manner similar to that of normal, staminate involucres in typical 
plants. The all-pistillate condition has been well known 
previously in a variant of A. artemisiifolia, which, like the fore- 
going, bears no staminate flowers at all (Jones, 1936, fig. 6), 
and which is found with the normal type and intermediates 
throughout Michigan. The all-pistillate condition is unknown, 
however, in A. coronopifolia, which, so far as is known, always 
comprises plants of the normal, bisexual type with axillary 
pistillate flowers and terminal racemes of staminate flower 
clusters. It seems entirely possible, therefore, that the pistillate 
intermediate population at Interlochen arose as hybrid in which 
the female parent was the pistillate form of A. artemisiifolia. 
Jones (1943) succeeded in crossing another ragweed, A. trifida, 
which is, like A. coronopifolia, a strictly monoecious species, with 
the pistillate form of A. artemisiifolia and showed that the 
progeny contained, in addition to monoecious and intergrading 
forms, the all-pistillate form as well. 

In 1910, Rydberg described a plant, Ambrosia media, as 
resembling A. coronopifolia in leaf shape. He wrote that 
"Otherwise, the plant is more closely related to A. elatior and 
A. artemisiifolia [which he regarded as separate species], the root 
being annual and the fruit spiny." Our examination of the type 
specimen of A. media, which was lent to us through the courtesy 
of Dr. Keck of the New York Botanical Garden, shows that the 
plant in question is the coarsely lobed form of A. artemisiifolia, 
a form not uncommon as a variant in the populations of this 
species in southern Michigan. S. F. Blake reduced this plant to 
synonymy under A. artemisiifolia (1925). We further checked 


196 Rhodora [Vor. 60 


the pollen grains of the type specimen and found that the average 
pollen size is 19.0 microns, the total range from 14.7 to 21.0, and 
the standard deviation 0.95. The good grains in a sample of 
1009 grains formed 79% of the total. Comparison with Table 1 
will show that these figures are well within the ranges charac- 
teristic of A. artemisiifolia and not of A. X intergradiens. 

FRUITS: For comparison of mature fruits of Ambrosia X 
intergradiens and its presumed parents, a field study was made 
on September 28, 1957, north of Stanwood, Mecosta County. 
In the prairie-like, rolling fields and roadsides there, all three 
taxa are common, and are readily compared as their fruits are 
fully developed at this season. In general, those of the parents 
may be immediately differentiated: those of A. artemisiifolia 
have terminal beaks twice as long on the average (1.2 mm.) as 
those of A. coronopifolia (av. 0.6 mm.). The lateral processes or 
“spines” of A. artemisiifolia average 0.5 mm. in length, while 
those of A. coronopifolia average 0.2 mm. and are commonly 
absent altogether in the latter as shown in Figure 3, C. There 
is, however, some variation and overlap as the specimens figured 
reveal. Any large collection of fruits of A. artemisiifolia in 
Michigan will show some individuals with very short lateral 
processes; and, as described earlier, a variant of A. coronopifolia 
exists (Yuma, Wexford Co., 8497a) in which the processes on 
the fruits are unusually well developed. Ambrosia X intergra- 
diens is intermediate between the two other taxa in fruit mor- 
phology (see Table I) and the rather well developed beaks and 
spines provide a valuable additional character with which to 
distinguish the new taxon from A. coronopifolia in late summer 
and fall. 

The pistillate flowers of A. artemisiifolia tend to occur in 
clusters of five or six in the axils of the upper leaves, but those 
of A. coronopifolia tend to be solitary (as determined from an 
average of 100 pistillate inflorescences for each species). "The 
average number of flowers in inflorescences of A. X intergradiens 
is intermediate, 1.e., averaging three flowers. The percentage of 
fruits which actually enlarge and mature differs considerably in 
the three taxa: In A. artemisiifolia, a total of 541 pistillate 
flowers counted yielded 415 approximately full-sized fruits, a 
proportion of 77%. In the plants identified as A. coronopifolia, 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 197 


a total of 118 flowers formed only 66 fruits, i.e., 55925. In the 
hybrid taxon it was very low—of the 280 flowers counted, only 
34 had expanded into normal-appearing fruits, only 16%. 
Ambrosia X intergradiens thus shows morphologically a high 
degree of sterility, a condition suggested also by the percentage 
of bad pollen grains and by the irregular meiotic process to be 
described below. 

It is interesting to note that the clone of wholly pistillate 
plants of A. X intergradiens discovered at Interlochen, Grand 
Traverse Co., failed entirely to set fruits, and repeated collections 
made during September and October, 1957, of many pistillate 
spikes revealed not a single fully formed fruit. 

POLLEN GRAINS: Following the same technique described above, 
the average pollen diameter of 12 collections of A. X intergradiens 
was determined as 21.8 microns, the total range of individual 
grains from 16.6 to 29.6. That the pollen grains of the inter- 
mediate ragweed turned out to average larger than those of A. 
coronopifolia was surprising, in view of their respective chromo- 
some complements which would lead one to expect the reverse 
relationship. However, the variation in size of the pollen grains 
of the intermediate proved to be considerably greater than that 
of either of the parental ragweeds, including A. coronopifolia, 
and the standard deviation was 2.30 microns for the former and 
1.05 for the latter. 

Although two of the collections of A. coronopifolia revealed a 
high percentage of non-staining pollen grains (one with 33% and 
one with 72%), most showed a low percentage and the average 
of 11 collections was 22.6%. On the contrary, all of the collec- 
tions of A. X intergradiens had high percentages of abortive 
grains. The mean of all the collections examined was 54.9% 
bad grains, the lowest single collection 424 and the highest 83 Z. 
These data on abortive pollen and pollen diameters are sum- 
marized in Table I, along with similar facts concerning A. 
artemistifolia for comparison. 


* If our figures are at all representative for the species as a whole, then A. artemisiifolia 
with its much larger number of pistillate flowers and greater average production of fruit per 
head has well over six times the reproductive potential by seeds as A. coronopifolia. Actually 
the difference is probably even greater because of the tendency for more axils to form pistillate 
inflorescences in A. artemisiifolia, Thus the annual species which relies entirely on seeds for 
survival, (so far as we know) shows a striking difference in seed production from the perennial 
ragweed which relies on its ability to remain alive from year to year and its ability to propagate 
itself by underground roots. 


198 Rhodora (Vou. 60 


CHROMOSOMES: Meiosis in the intermediate ragweed shows 
conspicuous irregularities. First metaphase is characterized by 
univalents, bivalents, and trivalents, and first anaphase com- 
monly shows lagging of chromosomes. The determination of 
units from ten well-spread sporocytes from five collections 
averaged 14.5 univalents (range: 8-19), 15.1 bivalents (range: 
11-20), and 2.5 trivalents (0-4). The average total number of 
units was 32.1 (range: 22-38). A rough explanation for the 
average pairing behavior might be suggested as follows: There 
are 54 chromosomes present, 18 from A. artemisiifolia and 36 
from A. coronopifolia. Assuming a fairly high degree of ho- 
mology between chromosomes of the parents, it then appears 
that 15 of the chromosomes of A. artemisiifolia and 15 of A. 
coronopifolia tend, on the average, to form pairs. The remaining 
three chromosomes of A. artemisiifolia would form trivalent 
configurations with six chromosomes of A. coronopifolia, leaving 
a residue of 15 univalents of the latter. The situation, however, 
is very much more variable from cell to cell than such an 
idealized *average" behavior would indicate. 

The lagging that commonly occurs in the chromosomes of 
first anaphase result in the exclusion of up to as many as four 
chromosomes from the second metaphase division figures, so 
that they lie off the respective equatorial planes, as shown in 
figure 4, 9. At second anaphase, as many as ten chromosomes 
have been seen lying separate in the cytoplasm, and even after 
the nuclear membrane is formed, chromosomes may remain un- 
assimilated. As the pollen grains mature, these chromosomes 
apparently disappear. In material of the intermediate plant 
from Pellston, Emmet Co., a sample of 171 division figures 
showing second anaphase had 55% of the figures with excluded 
chromosomes. The remaining 45% appeared to have normal 
second anaphase figures’. In contrast, in material of A. coro- 
nopifolia taken at the same time and place, a sample of 39 
figures showed all normal second anaphases. 

Root tip squashes from three localities of the intermediate 
ragweed confirm 2n = 54 chromosomes, as would be expected 

6 Avers, C, J. (Genetics 39: 117-126, 1953) has suggested a mechanism in Aster whereby 
triploids produce fertile pollen by means of a double equational division of the univalents. 


No evidence of such a mechanism has been observed in the present study, but this is a possible 
explanation of the relatively high percentage of normal-appearing second anaphase figures. 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 199 


from the known genomes of the putative parents, A. artemistifolia 
with 2n — 36 (Jones 1943; Yuasa 1956), and A. coronopifolia 
with 2n = 72. These localities were in Benzie Co. (8335); 
Crawford Co. (8348); and Alger Co. (8438-2). Irregular meiotic 
behavior was observed in the Benzie Co. and Crawford Co. 
materials, as well as Osceola Co. (8429), Emmet Co. (8445), 
and other Crawford Co. collections (8448). 

DISCUSSION: As botanists did not find Ambrosia coronopifolia 
in Michigan until 1900 we may assume that perennial ragweeds 
were probably introduced into the state sometime prior to the 
turn of the century. It should be pointed out, however, that the 
question of “introduced” vs. "indigenous" may be a vexing one 
when weeds are involved. Our concepts of these terms do not 
ordinarily admit of degrees (such as "introduced in part," or 
“mainly introduced"). With regard to our perennial ragweeds, 
it seems entirely plausible that from time to time, over thousands 
of years, small ''extra-territorial  emigrations from the home 
range into Michigan took place. Many, if not all, of these 
probably died out. However, now— with railroad and highway 
systems, and myriad artificially disturbed habitats, the species 
'an readily migrate into Michigan along definite pathways from 
the states further west. So even if the species had been present, 
though extremely rare, prior to 1900, the bulk of present popula- 
tions may very well have immigrated from further west. 
Certainly over its present range in the state, the perennial rag- 
weed will have to be interpreted as “mainly introduced." There 
is no positive evidence to date, in faet, to indieate that 1t was 
ever native at all. 

When Michigan in its entirety is considered, A. coronopifolia 
must be treated as only frequent to uncommon; but its ability to 
"take over” large areas locally where the habitat is appropriate 
make it nevertheless a potential hayfever problem in its restricted 
territories. Its habit of congregating its populations around 
settlements make its hayfever significance out of proportion to 
its over-all abundance. The annual species, A. artemisiifolia, 
will in general much outweigh its allergic importance because of 
greater numbers. But in the two weeks prior to flowering in 
annual ragweed, the perennial species may become a local 
problem to allergic persons. 


200 Rhodor: [Vor. 60 


The hybrid perennial ragweed has turned out to be remarkably 
common in appropriate localities. If it had been found only 
once or several times in this investigation it would have been 
designated only by formula. The intermediate has been given 
a taxonomic binomial, Ambrosia X intergradiens, because of a 
belief that any natural hybrid which comes to form many indi- 
viduals of importance in the community, whether by sexual 
means (e.g., as allopolyploids) or by asexual means (e.g., by 
underground reproductive axes, as in this instance) or both, 
should be so named. Perhaps other ragweed hybrids (e.g., A. X 
helenae Rouleau, an extremely rare plant in Michigan) are 
formed de novo in nature as frequently as A. X intergradiens, 
but none of the others, to our knowledge, compare in abundance 
to the present one which is self-perpetuating and builds up large 
local populations through the years. 

Now that A. X intergradiens has been distinguished among 
the populations of perennial ragweeds in Michigan, collectors 
may be urged to look for it in other states (e.g., Illinois, Minne- 
sota, and Wisconsin) where the two parental species intermingle. 
The characteristics of the new ragweed are subtle ones, it is true, 
and they are hard to perceive on easual inspection in the field; 
but the description and figures given above should suffice for its 
recognition. 

The sizes of pollen grains reported here for the plant commonly 
referred to as **western ragweed" in the Great Lakes states differ 
from the previous reports. Wodehouse (1928, 1945) gave the 
diameter of pollen grains of “A. psilostachya” as 23.4 (22.0-27.4) 
microns. His measurements evidently refer not to our plant but 
to other taxa in the complex. Pollen grains in A. coronopifolia 
as defined here measure 20.5 microns in diameter on the average. 
Our own measurements of A. psilostachya from California 
(Solano Co., Heiser 1966; San Diego Co., Alderson s.n.; Colusa 
Co., Chandler s.n.; Stanislaus Co., Hoover 165; Los Angeles Co., 
Wolf 4241; and San Luis Obispo Co., Summers s.n. —MICH and 
UC) gave an average diameter of 23.0 microns, the range 20.0— 
25.0. There is a correlation of pollen grain diameter with 
chromosome numbers. Previous studies of chromosomes in the 
genus Ambrosia indicate that the X number is 12, 17, or 18 
(Darlington & Wylie, 1955). Jones (1933, 1943) showed the 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 201 


chromosome number in A. artemisiifolia (as A. elatior) to be 
n = 18;in A. bidentata, n = 17; and in A. trifida, n = 12. Of 
these species the first is most obviously related to the perennial 
forms under discussion here, and the number of » = 36 de- 
termined by us (1957) and confirmed by Mulligan (1957) for 
A. coronopifolia supports this relationship. Ambrosia corono- 
pifolia may therefore be considered a tetraploid species. Ambro- 
sia X inlergradiens would then be a triploid. Heiser and 
Whitaker’s (1948) report of California material of A. psilostachya 
(Solano Co., Heiser 1966, UC) as having an estimated “n = 
50—52" suggested to us that the actual figure might be n = 54, 
i.e., the hexaploid number. Materials kindly collected for us by 
Mrs. D. A. Beals in Los Angeles Co., California (MICH) were 
observed in Metaphase I and Metaphase II: a total of 20 esti- 
mates ranged from 50 to 56, the average n = 53. The exact 
number may, accordingly, really be n = 54. It is worthy of 
mention at this point that smaller-spored forms also occur in the 
perennial ragweeds of California, suggesting that tetraploid taxa 
exist there as well as hexaploid. "Three collections (San Diego 
Co., Palmer 161; Lathrop, Walker 889; and Yuba Co., Howell 
28288—all UC) averaged 20.8 microns in diameter of pollen 
grains, with a range from 18.5-23.3, corresponding closely to the 
figures given above for A. coronopifolia. 

On the basis of present knowledge, therefore, we may assume 
that there are probably at least four polyploid levels in the 
A. artemisiifolia-coronopifolia complex, viz. 2x (artemisiifolia), 
3x (intergradiens), 4x (coronopifolia plus this or an additional 
taxon that grows in California), and 6x (psilostachya). Where 
it has been possible to compare them, the levels of polyploidy 
are matched by corresponding average pollen diameters, viz. 
17.6 microns (artemisiifolia), 20.5 microns (coronopifolia), and 
23.0 microns (psilostachya). Ambrosia X intergradiens does not 
conform with the sequence, but its irregular meiotie conditions 
and degree of spore abortion are such that the normal process of 
pollen ontogeny may be disturbed and thus produce the deviation. 

It can be concluded that polyploidy has played an important 
role in the evolution of the artemisiifolia-coronopifolia complex of 
ragweeds. Further knowledge of the evolution of this assemblage 
will have to be enriched by a survey much broader in scope 


202 Rhodora [Vor. 60 


than the present one. In particular, areas of presumed geo- 
graphical origin should be sought and investigated. Professor 
G. L. Stebbins has written. (December 18, 1956) that in his 
opinion the understanding of the relationships of the taxa in this 
complex will require “a careful study of all the forms occurring 
in the southwest United States, Mexico, and elsewhere in the 
American tropics.” We might suggest then that the forms that 
are found in California and in Michigan probably represent 
peripheral end-point populations derived from an evolutionary 
matrix that originated in the south. An interesting further 
possibility was recently indicated by Yuasa (1956) when he 
pointed out that "since the basic number of chromosomes in 
Compositae 1s nine, it is not surprising that the chromosome 
number of the ragweed [i.e., A. artemisiifolia] is n = 18.” If 
the original number in these plants was x = 9, then the common 
annual ragweed is a tetraploid species itself; A. coronopifolia 
would have to be interpreted as 8x; A. X <intergradiens 62; 
and A. psilostachya 12x. 
SUMMARY 

1. A study of perennial ragweed populations in the state of Michigan 
was made from standpoints of their distribution, periodicity, habitats, 
morphological variations, chromosome numbers, and pollen sizes. 

2. The known Michigan range of Ambrosia coronopifolia was increased 
by 18 new counties in field studies, bringing the total to 43. The species 
was found to be frequent across the Upper Peninsula to the north half of 
the Lower Peninsula and the western part of the southern half. Historical 
evidence does not favor interpreting it as originally native; in faet, no 
colleetions are known prior to 1900. 

3. It forms large clones by proliferation from underground parts in 
disturbed habitats such as roadsides and railways, especially around 
populated areas. Unlike the annual species (A. artemisiifolia) the 
perennial will invade grassy fields. The substrate is normally sterile, 
sandy or gravelly soil, in exposed places. 

4. Ambrosia coronopifolia matures earlier than the annual species by 
two or three weeks as evidenced by gross morphology and pollen pro- 
duction. 

5. Its variations include those of the leaves (lanceolate to deltoid in 
outline, from coarsely lobed to bipinnatifid, and sessile to short-petiolate), 
of leaf arrangement (opposite, whorled, and alternate), of plant habit 
(small and simple, to large and ‘“‘bushy’’), and of spines on the fruit 
(from unarmed to moderately spiny). 

6. Its pollen grains are smaller (20.5 microns in diam.) than previously 
reported in perennial ragweeds, and an average of 23 per cent of those 


1958] Wagner and Beals,—Perennial Ragweeds (Ambrosia) 203 


tested lacked protoplasts. Its chromosomes number 2n = 72, and 
melosis appears to be normal. 

7. An heretofore undescribed perennial ragweed, A. X intergradiens, 
hybr. nov., differs from A. coronopifolia in hairiness, color, leaf cutting, 
petiole length, and fruit structure. Its characteristics are intermediate 
between the latter and the annual A. artemisiifolia and it is interpreted 
as their natural hybrid. 

8. The new ragweed is unexpectedly common and a large number of 
populations have been observed in 15 counties. It grows usually, though 
not always, in company with the parents. 

9. The primary variation of A. X intergradiens includes characters 
of hairiness and plant size. One peculiar sterile population was found 
with wholly pistillate flowers. 

10. The pistillate inflorescences average 3 flowers per axil and are thus 
intermediate between the parents. However, only 16 per cent of the 
flowers matured normal-appearing fruits, indicating low fertility. 

11. Pollen grain diameter of A. x intergradiens is greater (21.8 microns) 
than either parent, but the variation is twice that of the parents. Over 
half of the grains appear to be inviable. The chromosomes number 
2n = 54, but meiosis is irregular with many univalents and a few tri- 
valents at Metaphase I, and with a large number of figures showing 
lagging of chromosomes. 

12. Pollen size of various ragweeds is discussed in relation to polyploid 
levels: there is a direct correlation. Ambrosia artemisiifolia is interpreted 
as a diploid, A. coronopifolia a tetraploid, and A. psilostachya, a hexaploid, 
based on x = 18. Ambrosia x intergradiens is a triploid, but its pollen 
size does not conform, probably because of irregularities due to its im- 
mediate hybrid origin. 

13. A really comprehensive knowledge of this complex must entail 
studies in warmer regions in southern U. S. and Mexico. The primitive 
members of the complex may have n = 9 chromosomes. 


LITERATURE CITED 

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Bucunuorrz, K. P. et al. 1954. Weeds of the North Central States. 
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DawNiELs, F. P. 1904. The flora of the vicinity of Manistee, Mich. 
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DanLiNGTON, C. D. AND A. P. Wyte. 1955. Chromosome Atlas of 
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Dram, C. C. 1940. Flora of Indiana. Dept. of Conservation. Indian- 
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Doper, C. K. 1899. Flora of St. Clair County, Michigan. Ann. Rept. 
Mich. Hort. Soc. 

—— —. 1918. Contributions to the Botany of Michigan, II. 
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204 Rhbodora [Vor. 60 


——————, 1921. Miscellaneous papers on the Botany of Michigan. 
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FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American 
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GLEARSON, H. A. 1952. Illustrated flora of the northeastern United 
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Heiser, C. B. anb T. W. WHITAKER. 1948. Chromosome number, 
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Jones, K. L. 1933. Ambrosia bidentata Michx. X A. trifida L. Amer. 
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——————, 1936. Studies on Ambrosia. I. The inheritence of floral 
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699. 

— ———-,. 1943. Studies on Ambrosia: III. Pistillate Ambrosia 
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Kenoyer, Lesie A. 1934. Forest distribution in 8. W. Michigan as 
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Moss, E. G. 1956. Ragweed in southeastern Alberta. Canadian 
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MvuLuGaN, G. A. 1957. Chromosome numbers of Canadian weeds. 
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1958] Steyermark,— Botanical Area in Missouri 205 


AN UNUSUAL BOTANICAL AREA IN MISSOURI 
JULIAN A. STEYERMARK 


Since the time of John Bradbury in 1809, and subsequently, 
through the efforts of Engelmann, Broadhead, Swallow, Trelease, 
Letterman, Eggert, Kellogg, and others of that period, Missouri 
has received its share of botanical exploration. The later and 
more detailed explorations of B. F. Bush and E. J. Palmer in 
the late nineteenth century and the first three decades of the 
twentieth century revealed the existence of additional significant 
plant regions in the state. "Their work brought to light the 
coastal plain flora of the Mississippi Embayment of the south- 
eastern Missouri lowlands, the elements of a Great Plains flora 
in the loess hills of northwestern Missouri, the edaphic and 
special flora of the Grand Falls chert barrens, the southwestern 
floral element of the limestone “bald knobs" and southern floral 
element of the White River region of southwestern Missouri, the 
northern element in the flora of northern Missouri, and such 
relict floras as that of Jam-up Bluff in Shannon County. Thus, 
after nearly a century and a quarter of botanical endeavor, the 
possibilities of discovering unknown and significantly interesting 
floristic areas in the state would have seemed negligible. 

Nevertheless, during the past quarter century in a cycle of 
intensive exploration of areas, mostly remote and inaccessible to 
previous workers, the writer has revealed the presence within the 
state of such fascinating botanically rich areas as the sink-hole 
ponds and swampy meadows of the southeastern Ozarks, saline 
and brackish springs of central Missouri, and what represents a 
"driftless" area of northeastern and east-central Missouri. Each 
of these areas has uncovered many species new to the flora of 
the state, the results of which have, for the most part, appeared 
in various numbers of Rhodora. 

Within the past four years, a new area, previously unbotanized, 
but occurring in a section of the state thought to have been 
adequately explored, has been visited by the writer. Since six 
species new to Missouri have turned up in this area, it has been 
thought appropriate to devote a special article to the locality 
concerned, rather than to scatter the information in a more 
general report. 


206 Rhodora [Vor. 60 


The area concerned involves a series of sand hills and spring- 
fed streams emanating from Crowley Ridge in Stoddard County 
in extreme southeastern Missouri. These sands are of Tertiary 
age. Although they are also found in adjacent Scott, New 
Madrid, and Dunklin counties, the series developed in Stoddard 
County are larger and contain a more unusual flora than in the 
other sites. The particular area lies between three and one-half 
to three and three-quarter miles southeast of Bloomfield. The 
hills are generally forested with several species of oak (Quercus 
velutina, Q. coccinea, Q. imbricaria, Q. falcata), hickory (Carya 
texana, C. ovalis, C. tomentosa), Sassafras albidum, and Cornus 
florida, with an herbaceous forest floor cover of Panicum laxi- 
florum, Comandra Richardsiana, Arabis missiouriensis, Ascyrum 
hypericoides var. oblongifolium, Phlox pilosa, Lithospermum 
caroliniense, and Linaria canadensis. In the more open sandy 
areas such herbaceous plants as Polygonella americana, Euphorbia 
corollata, and Asclepias verticillata are common. In the small 
valleys along spring-fed streamlets between the sand hills occur 
stands of Ilex opaca of sizeable extent. Occurring with the holly 
are such shrubs as Alnus serrulata, Lindera Benzoin, and [tea 
virginica. Along the wet banks and swampy ground are found 
Osmunda cinnamomea, O. regalis var. spectabilis, Onoclea sensibilis, 
Dryopteris Thelypteris var. pubescens, rarely Woodwardia areolata, 
Panicum microcarpon, P. polyanthes, P. commutatum, Cyperus 
flavescens var. poaeformis, Scirpus rubricosus, Fimbristylis 
autumnalis, Carex virescens and C. lurida, Juncus diffusissimus, 
Habenaria clavellata and the rare H. ciliaris, Boehmeria cylindrica, 
Hypericum tubulosum var. Walteri, Jussiaea decurrens, Rhexia 
virginica, Chelone glabra, Solidago patula, S. rugosa var. cellidi- 
folia, the rare Eupatorium fistulosum, Pluchea petiolata, and the 
rare Polygonum arifolium. On adjacent dry sandy slopes the 
very rarely collected Aristida lanosa is found. 

Beginning the exploration of the area in August, 1954, the 
author found Bartonia paniculata at the head of a tiny spring 
branch and Pyrus melanocarpa along the bordering sandy banks 
in alder thickets and adjacent lower sandy slopes nearby. Both 
species were reported subsequently as new to the state (Rhodora 
57: 314, 315. 1955). The same year Paspalum setaceum was 
added from the same area, and in April, 1955, along the same 


1958] Steyermark,— Botanical Area in Missouri 207 


spring branch was found Trisetum pensylvanicum, also new. In 
June, 1955, Typha latifolia f. ambigua, was found as an addition 
along the same spring branch. Before having had an opportunity 
to report these latter discoveries, a visit to the area in October, 
1955, yielded Scleria nitida and Trichostema setaceum. 

The following is a summary of the data for the species from this 
unique area not previously reported for the state: 


Typha latifolia L., forma ambigua (Sonder) Kronf. Swampy 
spring branch at base of Crowley Ridge near junction with 
floodplain, on property of Mr. Martin, T 25 N, R 11 E, sect. 6, 
near Pleasant Valley Church and Triplett Cemetery, 5 mi. 
northeast of Dexter, Stoddard Co., June 24, 1955, Swyermark 
78660. This form was growing with typical T. latifolia (Steyer- 
mark 78661). 

Trisetum pensylvanicum (L.) Beauv. Along spring-fed 
creek near junction of Crowley Ridge and lowland, bordered by 
sandy wooded ravine slopes, T 25 N, R 11 E, NW 14 sect. 6, on 
property of Mr. Martin, 3l5 mi. southeast of Bloomfield, 
Stoddard Co., April 25, 1955, Ste ermark 78285. The author 
first collected this species in April, 1955. Upon returning to the 
same locality two months later in June to show Drs. C. L. 
Kucera and Robert McDermott of the Botany Department of 
the University of Missouri the area concerned, no trace of this 
vernal-flowering species was evident. The plants had been 
growing half-submerged in the water of the spring branch among 
the thickets of Alnus serrulata. In the shallow water of the 
same stream and occurring with Trisetum pensylvanicum were 
numerous individuals of an unusual aquatic fungus with fleshy 
whitish stipe and dull orange sporangia. It was identified by 
Dr. Francis Drouet as Mitrula phalloides. The range of the 
Trisetum, according to Gray's Manual, is “Mass. to O., s. to 
Fla., Ala., and La." Apparently, this record represents the 
northwesterly limit of the species. 

Paspalum setaceum Michx. (typical). Dry sandy wooded 
slopes bordering ravines along and tributary to spring-fed creek 
near junction of Crowley Ridge and lowland, on property of 
Mr. Martin, T 25 N, R 11 E, NW 14 sect. 6, 315 mi. southeast 


á 


of Bloomfield, Stoddard Co., August 20, 1954, Steyermark 76726. 


208 Rhodora [Vor. 60 


Previously known from “Fla. to Tex. n. to SE Mass., L. IL, O. 
and Ky. and Mex.", according to Gray’s Manual, this is another 
one of the coastal plain species which extends into this section 
of southeastern Missouri. 

Scleria nitida Willd. Sandy and mossy slopes above and 
along spring branch in valley at Junetion of Crowley Ridge and 
bottomland, on property of Mr. Martin, T 25 N, R 11 E, sect. 
6, 3°4 mi. southeast of Bloomfield, Stoddard Co., October 17, 
1955, Steyermark 80375. Although this species is not recognized 
by Core in his revision of Scleria nor by Gleason in the New 
Illustrated Flora as distinct, and included by them under S. 
triglomerata, S. nitida appears amply distinet in having larger 
achenes and a close dense puberulence on the inner sheath of the 
leaf blades. The habitat of S. nitida in Missouri is also quite 
different from that of S. triglomerata. The latter occurs in the 
state on rocky limestone glades and prairies on soils of neutral 
to alkaline reaction, whereas S. nitida is found in the Stoddard 
County locality on sandy acid soil. It represents a remarkable 
extension of range of a predominantly southeastern species. 

Trichostema setaceum Houtt. Sandy openings on slopes of 
blowout of sandy prairie and slopes on east side of road and 
eroded gully, Crowley Ridge, T 25 N, R 11 E, sect. 31, 3!5 mi. 
southeast of Bloomfield, Stoddard Co., October 17, 1955, Steyer- 
mark 80349. Occurring in the same immediate area of the open 
sand slope as Paspalum setaceum, the discovery of this species 
adds another of the increasing numbers of coastal plain species 
to the flora of Missouri. Its previous known range was “Fla. to 
La., n. to sw. Ct., centr. Pa. and s. O.", according to Gray’s 
manual. The new station represents its most northwesterly 
known occurrence. It was associated in the open sand area with 
Aristida lanosa, Digitaria filiformis var. villosa, and Froelichia 
floridana var. campestris. 


MISSOURI BOTANICAL GARDEN. 


Volume 60, No. 714, including pages 145-176, was issued 11 July, 1958. 


ARLOW REFERENCE LIBRARY NOV 3 1958 


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Vol. 60 August, 1958 No. 716 
CONTENTS: 


Notes on Nearctic Hepaticae VI. Phytogeographical Relation- 
ships of Critical Species in Minnesota and Adjacent Areas of 
Great Lakes. R. M Schuster. 650.5. «i cewek ee 209 


Some New Floras for Parts of North America. R. C. Rollins 
(Review) ccc cae eee ees ee ve ehs s od on D MN 235 


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Vol. 60 August, 1958 No. 716 


Notes ON Nearctic Heraticaer VI 
PHYTOGEOGRAPHICAL RELATIONSHIPS OF CRITICAL SPECIES IN 
MINNESOTA AND ADJACENT AREAS OF THE GREAT LAKES 


R. M. SCHUSTER! 


A number of highly disjunct species of liverworts have recently 
been reported from the Great Lakes region by the writer in a 
manual on the Hepaticae of Minnesota, published in 1953. 
Limitations of space, and orientation of objective then prevented 
as full a discussion of the implications of some of the distribution 
patterns as would have been desirable. The present publication 
is intended to discuss the phytogeographical aspects which were 
not previously treated in detail. In utilizing the present paper, 
it may prove helpful to refer to the more generalized and abbre- 
viated discussion of the floristic affinities of the Hepaticae of 
Minnesota (Schuster, 1958). In that paper are given a series of 
maps which, in part, supplement the maps included here. 

The distribution patterns of the bryophytes have not received 
the attention that they deserve. This is partly due to the fact 

1 The field work leading to the preparation of this paper was supported in part by two grants 
from the National Science Foundation (NSF Grants 669, 1369); during a portion of the writing 
of the paper the writer held a Fellowship from the Guggenheim Foundation. I would like to 
express my appreciation to Drs. W. D. Billings (Duke University), Warren H. Wagner Jr. and 
Edward Voss (University of Michigan) for critically reading portions of this paper. In the 
preparation of the original drafts I was assisted by my wife, Olga M. Schuster, who has also 
assisted in preparing the final draft for publication. 

It should be pointed out that the original draft of this paper was written more than five years 
ago. 'The accumulation of additional phytogeographical data has been so rapid that the paper 
has been revised almost annually, greatly delaying its appearance. Several papers in this 
series (Nearctic Hepaticae, VIII-XI) have, in the meantime, appeared. The phytogeographical 
data is as complete as possible, up to, and usually including data published in 1956. I have 
omitted questionable reports, while localities that could not be accurately pin-pointed are 
indicated with a question mark on the maps. In a few cases stations are known for some 


species lying in the Aleutian chain of islands, beyond the margins of the map commonly used. 
An arrow is used to indicate a station (or stations) lying west of the area given on the map. 


210 Rhodor: | VoL. 60 


that an idea still persists that bryophytes do not show as rigidly 
defined ranges as the higher plants, and thus approach many 
algae in their distributional peculiarities, or lack thereof? The 
rate of evolution of the bryophytes is probably much slower than 
that of many of the genera and families of the angiosperms, with 
the consequence that there is a slower, and less marked tendency 
for disintegration into geographical races or varieties. Such 
“conservatism” of the Bryophyta is, however, of marked advan- 
tage for the student of plant distribution, in that the bryophytes 
may often serve to show past phytogeographical links which 
otherwise have nearly been destroyed. An example, if one is 
needed, is the established connection between the flora of western 
England and Ireland, with that of the Southern Appalachians, 
that of the coastal portions of Alaska, and that of eastern Asia. 
The existing distribution of Herberta sakurait serves to emphasize 
the floristic features these areas once had in common (Schuster, 
1957a). The phytogeographical links thus established probably 
represent remnants of a former, more widespread range. That 
such connections remain is perhaps partly the consequence of a 
high “somatic plasticity," and correlated lower evolutionary 
potential, of many species of Hepaticae. This is especially the 
case with species which reproduce largely or exclusively by 
asexual methods. In such cases the chance for genetic recom- 
bination is proportionally infrequent or even lacking. 

Linked with the inherent evolutionary “conservatism” of many 
bryophytes is their small size, and consequent ability to survive in 
limited niches, or microenvironments. The reduced size, as 
compared with that of seed plants, allows them to survive in 
small niches, even if they lack the ability to evolve to meet new 
environmental conditions. In a similar situation, the angio- 
sperms in most cases must either evolve and adapt, or else die out. 
As a result, bryophytes, in many cases, perhaps have remained as 

2 An equally significant factor in limiting the use of phytogeographical information derived 
from the Bryophyta is the difficulty of access to such data, and the relatively limited amount 
of such data available. The effect of recent collections on our concepts is illustrated, for 
example, by the ranges here plotted for Diplophyllum obtusifolium and Anastrophyllum saxicola. 
The writer has collected, in recent years, all of the material from eastern United States of both 
of these species, with the exception of only one collection. The recognition of these species as 
elements in the flora of the eastern United States thus is a very recent matter. Equally recent 
is our still very partial knowledge of the American range of Scapania scandica, which is not 


listed for the western hemisphere in the recent (1938) list of Buch. Evans & Verdoorn, and still 
listed as only known from Greenland in Frye & Clark (1946, in 1937-47). 


1958] Schuster,—Notes on Nearctic Hepaticae VII 211 


relicts in sites where almost all disjunct angiosperms have dis- 
appeared and their study, therefore, is pertinent to the whole 
problem of the evolution of our present flora. 

Within the more restricted framework of the phytogeographi- 
cal affinities of the Great Lakes region, the pertinence of phyto- 
geographical data, derived from bryophytes, has been shown in 
the pioneer paper of Steere (1937). He pointed out that the 
same phenomena of disjunction existed among the Great Lakes 
bryophytes as had been demonstrated by Fernald (1925, 1935) 
for the tracheophytes. In Steere’s paper, the disjunct Bryophyta 
were treated as members of four floristic elements, as follows: 


(1) Aretie species which reach their southernmost point in the Lake 
Superior region. | Asterella ludwigii (Fig. 7) is cited as an example. 

(2) Almost strictly Cordilleran species, "usually characteristic of high 
altitudes.” | Jungermannia schiffneri is cited as an example, and a map 
of its then-known distribution is given (Steere, Fig. 4). 

(3) Pacifie coast species, usually characteristic of lower altitudes. 
Frullania bolanderi is cited as an example, and its distribution given 
(Steere, Fig. 5; see Fig. 12). 

(4) Cordilleran species, also found (at lower elevations) in the ‘‘Driftless 
Area” of Wisconsin, Iowa, Minnesota and Illinois. An example is cited, 
Asterella saccata (see Fig. 14). 


To these distributional types the present writer would add, for 
the purposes of this discussion, several others: 

(5) Arctic species, occurring in the "Driftless Area," but evidently not 
in the Lake Superior Region. Athalamia hyalina (Fig. 15) and Mannia 
sibirica (Fig. 16) have distribution patterns of this typo. 

(6) Aretie species, occurring southward to the Lake Superior Region, 
and also in the “Driftless Area." — Mannia pilosa (Fig. 16) has a distribu- 
tion pattern of this type. 

(7) Appalachian species, with a scattered range into the Ozarks, in 
some cases to the "Driftless Area" and northward also to the Lake 
Superior region. Mannia rupestris and Diplophyllum apiculatum are 
examples (Figs. 17, 18). 

The total effect of the introduction of these additional more or 
less clearly disjunct vegetational elements serves to attribute to 
the Great Lakes Area (including the nearby ‘‘Driftless Area"), 
and specifically the region peripheral to Lake Superior, an ex- 
tremely synthetic and diversified flora, considering that the entire 
region appears to have been glaciated. In the following discus- 
sion, examples of each of the vegetational elements are treated, 


212 Rhodora [ Vor. 60 


followed by a discussion in which these discordant distribution 
patterns will be briefly analyzed.? 


(1) ARCTIC SPECIES, REACHING THEIR SOUTHERNMOST RANGE IN THE 
LAKE SUPERIOR REGIONS (Figs. 1-8) 


In general, the species with this type of distribution occur southward 
(at high elevations) to New England, and (at much higher elevations) 
southward to varying degrees in the Cordilleran chain, Exceptions are 
specifically noted. 

ODONTOSCHISMA MACOUNII (Aust.) Underw. (Fig. 1). This species, of 


T 
J 


Fig. 1. North American range of Odontoschisma macounii. 


arctic and alp' ne distribution in Europe and North America, occurs in a 
transcontinental belt, lying largely between 55°-82°21’ Lat. N. Eastward 
it occurs to the south only on Cape Breton I., and on islands in Lake 
Mistassini; westward it occurs to St. Matthew I. south of Bering Strait. 
In the United States, it is found only in a restricted area along the North 
Shore of Lake Superior from Minnesota to Ontario (type), and in Wis- 
consin and Michigan along the southern shore of Lake Superior. The 
range is somewhat restricted by the weakly “calciphilous” nature of the 
species. 

3 Subsequent to the completion of the text and maps, the important paper by Buch & Tuomi- 


koski (1955) on the Hepaticae of Newfoundland, has come to hand. Several of the species 
treated herein are there extended to Newfoundland. 


19581 Schuster,— Notes on Nearctic Hepaticae VII 213 


Fig. 2. North American range of Lophozia (Leiocolea) gillmani. 


LoPHOZIA GILLMANI (Aust.) Schuster (Fig. 2). A relatively widespread 
arctic-alpine species more strongly restrieted than the last by pronounced 
Ca requirements. Found in isolated sites, from northeastern Ellesmere I. 
(Schuster, 1955) south to Quebec, Nova Scotia, Vermont and New Hamp- 
shire (to the south at medium and high elevations), as well as in the west 
from the Yukon to the south coast of Alaska, to northern Oregon and 
Montana. Abundant in the Great Lakes area from Manitoulin I. in 
Lake Huron to Lake Superior. 

SCAPANIA MICROPHYLLA Wstf. (= S. buchii Müller) An arctic-alpine 
species known from northern Europe, westward to Iceland, reported from 
one station in Maine, and from Prince Edward I.; recently reported from 
the north shore of Lake Superior, Lake Region between Minnesota and 
Ontario, and from Isle Royale, Michigan (Schuster, 1953). An un- 
published report extends the species to Wisconsin. This systematically 
diffieult species probably will prove transcontinental when more fully 
understood. 

Lopuozia (Orthocaulis) QuapRILOBA (Ldb.) Evs. (Fig. 3). Usually 
considered a high arctic species, with the range in North America almost 
totally north of the Arctic Circle, north to 83°6’ N. in Greenland and 
82°32’ N. in Ellesmere I., only two reports from the high mountains of 
the Canadian Rockies lying far to the south. Recurring, totally as a 


214 Rhodora [ Vor. 60 


disjunct, at Copper Harbor, and at the Devil's Washtub," Keweenaw 
Co., Michigan (Schuster, 1953). 

Lornozia (Leiocolea) scnuurai (Nees) Schiffn. In Europe essentially 
arctic in distribution, but in North America with a peculiar range in- 
volving Alaska and the Yukon, the west coast of Hudson Bay (Schuster, 
1955), the Gaspé, and the Lake Superior-Lake Michigan region. The to- 
tally anomalous report of the species from Bergen Swamp, N. Y. (Schus- 
ter, 1949) must be interpreted in the light of the occurrence at the latter 
locality of many western and Great Lake: species, i.e., Solidago 
hougl.tonii ; the species there is certainly a late Pleistocene relict. 

In the absence of reports from the eastern Arctic, the range of this 
species, in North America, approaches the next group to be treated, the 
Cordilleran species. However, it must be kept in mind that the species 
is an obligatory caleiphyte. 

SCAPANIA SCANDICA (Arnell et Buch) Maev. (Fig. 4). Until recently, 
the only reports of this species from North America were those of Buch 
(1928) from Greenland. Persson (1946) reported the species first for 
North America proper, the species occurring in four widely separated 


Fig. 3. World range of Lophozia (Orthocaulis) quadriloba. Not all stations could be indicated 
in cases where individual stations are situated less than 100 miles apart. 


1958] Schuster,—Notes on Nearctic Hepaticae VII 215 


Fig. 5. North American range of Scapania gymnostomophila (including S. incurva); the 
British Columbia report uncertain. 


216 Rhodora [ VoL. 60 


localities from Sitka on the southern coast to Kodiak, to St. Matthew I., 
and to Unalaska (the latter station is not indicated on the map). Persson 
indicated a probable oceanic distribution for the species. From a study 
of the entire range of the species I postulated that it should be found 
southward to Labrador, Newfoundland and Nova Scotia, in the same 
general areas as Diplophyllum albicans. Shortly thereafter, two collections 
were made by the writer on Mt. Katahdin, Maine: one near the foot of 
Hamlin Ridge, at only 2800 ft.; the other near the top of the Saddle 
Slide, at ca. 4500-5000 ft. (Schuster 32910, 32994a, 1954). A year later 
the species was found in Newfoundland (Buch & Tuomikoski, 1955). 
Later during the same year, a specimen was studied from the Lake 
Superior region, Oak I., Wisconsin (Cheney 6033, 1896, as Scapania 
curta; NYBG) which exhibited many similarities to S. scandica, but some 
to the extremely closely allied S. helvetica, a species known only from the 
European Alps. It proved to be almost impossible to decide which of 
the two taxa was involved, so the material was submitted to the European 
sp. eialist on the genus, the late Dr. K. Müller. He expressed the opinion 
that it was S. scandica. Although the occurrence of the high-arctic 
Scapania spitzbergensis on Mt. Katahdin (Schuster, 1951a) makes the 
occurrence there of S. scandica a matter of little surprise, the occurrence 
of S. scandica in the Great Lakes region is remarkable. 

SCAPANIA GYMNOSTOMOPHILA (Kaal.) Kaal. (Fig. 5) (inel. Diplophyllum 
incurvum Kaal.). An arctic species in Europe, with isolated Alpine 
stations, and one or two in the intervening lowlands. In North Americ: 
known from numerous stations on Ellesmere I., north to 82°32’ N. and 
recently established as occurring in the Yukon (Persson, 1952), with a 
somewhat dubious report from British Columbia (based on a Brinkman 
collection) and a dubious report from northern Idaho (Frye and Clark, 
1946). Again found from Cape Breton I. and Nova Scotia to Maine 
and Vermont, and ina rather large number of stations in the Lake Superior 
area. The distribution strongly restricted by the presumably obligatory 
Ca requirement. 

SCAPANIA CUSPIDULIGERA (Nees) K. Müller (Fig. 6). A species with 
as rigid a restriction to calcareous rocks as S. gymnostomophila, but with 
a wider distribution in North America. In Europe, aretic-alpine. In 
North America widespread from Alaska to Ellesmere Island and Green- 
land, southward in the Cordilleras to Colorado and New Mexico, and 
probably to California (at high elevations only!); in the east south to the 
east shore of Hudson Bay (Schuster, 1951), and to the St. Lawrence 
Valley and the Gaspé. Recurring, not infrequently, along the Lake 
Superior shore in Minnesota, Michigan and Wisconsin. 

I have also seen a single collection, of typical material, from the alpine 
portions of Japan (Inoue). 

ASTERELLA LUDWIGI (Schwiigr.) Underw. (Fig. 7). Also a "calciphile' 
and consequently of restricted range. With the exception of a doubtful 
report from Alaska, restricted to the Cordilleran region of the west, 
(where frequent at high elevations), and recurring along both the east 


1958] Schuster,—Notes on Nearctic Hepaticae VI 217 


Fig. 6. North American range of Scapania cuspiduligera. The report of this species from 
western New York (fide W. C. Steere), cited in Schuster (1949) is regarded as questionable. 


and west coasts of Greenland, in Baffin I., and at Ungava Bay; a doubtful 
report from the Gaspé. Known from four stations in the Lake Superior 
region (Ontario, Minnesota and Michigan). The rapidly increasing 
knowledge regarding the distribution of this species is clear if the map 
here given is compared with that in Steere (1937, Fig. 2). It is also 
illuminating to compare the distribution given for the species by Steere 
(1937) and by Frye and Clark (1937, in 1937-47). The latter, due to an 
inadvertent confusion of both the distribution data and the illustrations 
of the two species (see Schuster, 1953) gave the impression that A. 
ludwigii was a common lowland plant in eastern North America. They 
"extended" the range south to Missouri and Nebraska, and were followed 
in this by Whittlake (1954), who was able to “extend” the range of this 
species even as far south as Arkansas. All these reports are certainly 
erroneous, and must refer to A. tenella, the common lowland species. 
ANTHELIA JURATZKANA (Limpr.) Trev. (Fig. S). A non-caleiphile, of 


218 Rhodora [ Vor. 60 


Fig. 7. North American range of Asterella ludwigii. The report from the Gaspé uncertain; 
all of the reports of this species from eastern United States in Frye & Clark (1937) are erroneous. 


arctic-alpine distribution, known from the southern coast of Alaska to 
the Alberta-British Columbia boundary region, south to Wash ngton, and 
California (locality uncertain), and in the east from Greenland and 
northernmost Ellesmere I., south to the east coast of Hudson Bay. With 
a somewhat isolated occurrence on the summit of Mt. Katahdin, Maine 
(Schuster, 1949), and in the White Mountains of New Hampshire. 
Totally disjunct at Pictured Rocks, Michigan, on the Lake Superior 
shore. 

CEPHALOZIA LEUCANTHA Spruce. A low-grade arctic and subarctic 
species, apparently lacking in Greenland and Ellesmere L, but with a 
widespread distribution in the southern portion of the Tundra, and the 
ecotone between Tundra and Taiga in Quebec, Labrador, end Newfound- 
land. In addition, a single report from Mt. Marey, New York (Schuster, 
1953), and one from Minnesota (Schuster, 1953). In the west apparently 
frequent from Alaska, as far west as Agattu I., southward to British 
Columbia and Washington. 


1958] Schuster,—Notes on Nearctic Hepaticae VIT 219 


ANASTROPHYLLUM SAXICOLA (Schrad.) Schuster (Fig. 9). A rare 
arctic-alpine species, first reported from the United States from Pigeon 
Point, Minnesota (Schuster, 1953) but found recently on Mt. Katahdin, 
Maine (Schuster, August, 1954). The species occurs seattered from 
Alaska (as far west as Attu I.; station not on map) and the Yukon, and 
doubtfully to British Columbia, to east Greenland, Baffin I. (Stephani; a 
recent collection studied from Pangnirtung, Wynne-Edwards), Quebec, 
and the shore of James Bay in Ontario. The species is an oxylophyte. 

Very recently a wholly disjunct station of A. saricola has been dis- 
covered by the author on Roan Mt., both in Tennessee and North Carolina. 


Fig. 8. North American range of A nthelia juratzkana. 


The occurrence of this arctic-alpine species in the Southern Appalachians, 
although at first very surprising, is not without parallel. Sedum rosea, 
with the bulk of its range equally in the treeless regions from Labrador to 
Greenland, occurs as a disjunct on Roan Mt. 

ODONTOSCHISMA ELONGATUM (Ldb.) Evs. An essentially aretic-alpine 
species, rare in the Taiga. Widespread in Alaska; recurring eastward 
on the Greenland coast, the coast of Hudson Bay (Schuster, 1951), 
southward to the high mountains of Maine and New Hampshire. Re- 
curring as a disjunct, in the Lake Superior area, near Thunder Bay, 
Ontario, and at Deer Lake, west of Munising, Michigan (Schuster, 1953). 
Strongly oxylophytie. 

CEPHALOZIELLA SPINIGERA (Lindb.) comb. n. (based on Cephalozia 
spinigera Lindberg, Musci scand. 4, 1879; C. subdentata Wstf., Krypt. Fl. 


220 Rhodora [ Vor. 60 


Fig. 9. North American range of Anastrophyllum (Eurylobus) saxicola. 


Mark Brandenburg 1: 234, 1902). This helophytie species was first 
reported from continental North America by Schuster (1949), from 
western New York—the least likely place in which it would be expected 
to occur, because of the pronouncedly arctic and high subarctic character 
of the plant. The bulk of its range extends from St. Lawrence I. and 
continental Alaska to northern Quebee and Greenland. Collected 
several times in the high mountains of New England by the writer (Mt. 
Monadnock and Zealand, N.H.; Mt. Katahdin, Maine), and recently 
from northeastern Minnesota (Schuster, 1953). The plant is also present 
in a collection of Microlepidozia setacea made in a bog near Burt Lake, 
Michigan (Patterson). This species will probably prove transcontinental 
in peat bogs and on peat-covered ledges, with the bulk of its distribution 
in the Arctic. 

SCAPANIA DEGENII Schiffn., ex K. Müll. (Fig. 10) and Scapania hyper- 
borea. ‘These closely allied species (Schuster, 1953) are treated as a unit 
here because of the lack of agreement as to whether the Great Lakes p'ants 
are to be regarded as S. hyperborea or S. degenii. Because of their pecu- 
liarities, the Lake Superior plants have been regarded as a variety (var. 
dubia) of S. degenii (Schuster, 1953). S. hyperborea s. str. is frequent, 
apparently, on the Greenland coast, and occurs on the eastern shore of 


1958] Schuster,—Notes on Nearctic Hepaticae VI 221 


Fig. 10. North American ranges of Scapania degenit, including var. dubia (rings) and S. 
hyperborea (dots). 


Hudson Bay (Lepage, 1953); it was earlier reported by Evans from 
Maine and New Hampshire, where it is limited to the alpine peaks of the 
higher mountains. The closely allied S. degenii occurs in northernmost 
Quebec, the Belcher Islands in Hudson Bay, the east coast of Hudson 
Bay, and near Churchill on the west coast of Hudson Bay, and then 
recurrs again on the north shore of Lake Superior in Minnesota (Schuster 
1951, 1953); a specimen has recently been seen from the coast of Labrador. 
It was reported from North America only recently, almost simultaneously 


222 Rhodora [ Vor. 60 


by Arnell (1950) and Schuster (1951), and does not appear in the work of 
Frye and Clark. In the opinion of the writer, two distinet species are 
involved, S. hyperborea, an oxylophyte; 5. degeni;, usually a "weak" 
calciphyte. 

The preceding examples could be further supplemented, with- 
out too much point. The ranges of an additional series of species 
with a similar range are mapped, although not discussed, in 
Schuster (1958). They demonstrate the existence, in the Lake 
Superior area, of a well-developed element of arctic and arctic- 
alpine species, which extend further southward only in some 
instances (and then at high elevations in the mountains, in all but 
isolated cases). This, in the opinion of the writer, is clearly due 
to a microclimatic tundra zone, forming a narrow fringe around 
the coldest of the Great Lakes. The preceding species, therefore, 
are to be regarded as a sort of “rear guard” of a larger series of 


Fig. 11. North American range of Jungermannia polaris (open areas and rings) and of the 
presumed synonym, J. schiffneri (dots). 


1958] Sehuster,——Notes on Nearctic Hepaticae VII 223 


arctic species, at one time presumed to have characterized much 
of the raw, recently uncovered soil and rock, at the margin of the 
receding Pleistocene icesheet. The fact that of these 15 species 
(haphazardly selected), 8 are to be classified as ''calciphiles" is 
clearly not contradictory to such an explanation. These species, 
then, are simply relicts, left behind during the northward march 
of the vegetation after the last glaciation. Their survival 
locally, in such large numbers, is clearly partially due to the 
influence of the cold waters of Lake Superior. 

Subsequent to the completion of this paper, the writer had 
occasion to spend a summer in northeastern Ellesmere Island 
some 40 miles south of the northern tip and 80 miles south of the 
northern tip of Greenland (the northernmost land area in the 
world), collecting between Latitudes 82° 24’ and 82° 32’ N. 
Of these 15 species six (Odontoschisma macounit, Lophozia 
gillmani, L. quadriloba, Scapania gymnostomophila, S. cuspiduli- 
gera, Anthelia juratzkana) were found in this area; all of these are 
“calciphiles” or at least strongly Ca-tolerant. Of the many other 
largely arctic-alpine species reported for the Lake Superior area, 
several others were found, among them: Cephaloziella arctica, 
Lophozia kunzeana, L. heterocolpa, Lophozia latifolia and Crypto- 
colea imbricata. The latter two species, only recently described, 
very rare and restricted in the Lake Superior area, proved to be 
quite frequent on the north shore of Ellesmere I.! 


(2) CORDILLERAN SPECIES OCCURRING AS DISJUNCTS IN THE 
LAKE SUPERIOR REGION 


Steere cites Jungermannia schiffneri Loitlesb. (Fig. 11) as an example 
and gives a map which appears to support this viewpoint. However, 
this species has recently been reported from Quebee (Lepage, 1945) and 
is doubtfully known from Ellesmere I., while dubious material is reported 
as this species from western Pennsylvania. It is probable that a spo- 
radically distributed, widespread arctic-alpine taxon is involved, rather 
than a bona-fide Cordilleran species. The rarity of the species makes it 
unwise to generalize as to its floristic affinities.* 

4A further complicating factor in interpreting the range of Jungermannia schiffneri is the 
status of the plant known as J. polaris. This last was described (as J. pumila var. polaris) 
from Spitzbergen material, distributed by Berggren, No. 176, in the Musci Spitzberg., in 1868. 
Two years later it was raised to a distinct species by Lindberg, and reported from Greenland. 
Bryhn (1906-1907) reported numerous stations of it from Ellesmere Is. Indeed, judging from 
the number of stations, this was, on the Second Thule Expedition, the most frequently collected 
member of the Jungermanniaceae. J. polaris was, however, placed as a synonym of the 
dioecious Jungermannia atrovirens by both Stephani and Müller. (Con’t. p. 224.) 


224 Rhodora [ Vor. 60 


Perhaps no other good case of a species of Hepaticae with a restriction 
to the Cordilleran area, and an extension to the Lake Superior area is 
known. This, it should be emphasized, is most easily explained because 
of the essential lack of a Cordilleran series of species among the Hepaticae. 
With the greater age of the species, there has been, in virtually no case, 
divergent evolution of the widespread northern species with an east-west 
distribution in North America. If maps 1-11 are studied (of the arctic- 
alpine species), it will be seen that occasional species "approach" the 
Cordilleran-Great Lakes-St. Lawrence Valley pattern so often emphasized 
by Fernald. This is the case with Lophozia gillmani (Fig. 2) and also 
Lophozia schultzii, These species, however, are obligatory calciphiles, 
and the disjunction in distribution may be due largely to a disjunction of 
suitable habitats. As a consequence, the writer cannot at present 
recognize a Cordilleran element in the hepatic flora of the Lake Superior 
shore. 

The absence of such a Cordilleran element is accompanied by another 
phenomenon: the frequent Cordilleran extension southward, at high 
elevations of species I would regard as arctic-alpine in nature. Scapania 
cuspiduligera (Fig. 6), Asterella ludwigii (Fig. 7) and Scapania hyperborea 
(Fig. 10) all show such a southward range in the Cordilleras. Any real 
east-west disjunetion appears to be between an oceanic, Pacific element, 
restricted largely to the Coastal Ranges, and an Appalachian element. 
Among such examples may be cited Bazzania denudata and Herberta 
sakuraii (synonyms are H. hutchinsiae and H. tenuis; see Schuster, 1957a). 
East-west disjunction, in the Hepaticae, thus appears to be a phenomenon 
going back to the early Cretaceous, when the last great submergence of 
the North American continent took place. "This explanation, if tenable, 
would go far to explain the restriction of a high incidence of ‘‘endemics”’ 
to the area peripheral to the mid-Cretaceous, Mesocordilleran geanticlinal 
of the westernmost portion of the continent, and to the Appalachian 
region to the east. "This interpretation would involve, therefore, a pre- 
Tertiary and at least late Mesozoic origin of some extant species of 
Hepaticae. 


In 1955 I found that the only frequent member of the Jungermanniaceae found on the north- 
eastern coast of Ellesmere I. was the paroecious plant identical in all important respects with 
Jungermannia schiffneri. Subsequently, the collection of Berggren from Spitzbergen, described 
as J. pumila var. polaris, alluded to above, was examined. This collection must be considered 
as the type of var. polaris, although it is much less clear that it can serve as the type of the 
species polaris. Careful examination has revealed several pertinent features previously over- 
looked: (a) the plants are paroecious; (b) the capsules have the median epidermal cells of the 
valves with 2-4 strong, nodular thickenings on each face of alternating longitudinal walls 
while those longitudinal walls that alternate with the aforementioned walls lack thickenings or 
bear 1, rarely 2 thickenings. The cells average 13-15 u wide. The paroecious inflorescence, as 
well as capsule-wall anatomy are clearly characteristic of J. schiffneri, not of J. atrovirens 
suggesting that J. polaris is identical with the later described J. schiffneri! Certainly the 
northern Ellesmere material that I collected in abundance in 1955 is an exact match for the 
type of var. polaris. As a consequence, J. polaris (= J. schiffneri) now acquires a high arctic 
range, with isolated stations in the Cordilleras and the Great Lakes region, southward in the 
east to the montane portions of Quebee (Fig. 11). The species thus clearly acquires a range 
similar to the arctic and arctic-alpine species previously considered. 


1958] Schuster,—Notes on Nearctic Hepaticae VI 225 


Among such species, with a possible origin before the middle Cretaceous 
are the two following species, both of which are widespread at lower 
elevations on the Pacific Coast. These species are today found both 
east and west of the mid-Cretaceous Rocky Mountain geosynclinal, in 
which the great Cretaceous sea existed. 


(3) PACIFIC COAST LOWLAND SPECIES RECURRING IN THE 
GREAT LAKES AREA 


A disjunction of this type is, at first glance, extremely surprising. The 
classical case is that of Frullania bolanderi Aust. (Fig. 12), previously 
cited by Steere (1937). The considerable expansion in known range in 
recent years is evident on comparing the map in Steere (Fig. 5) with 
Fig. 12. 


Fig. 12. North American range of Frullania bolanderi; the species is otherwise known only 
from the Pacific Coast of northern Asia. 


F. bolanderi was described from near San Francisco, California. It 
extends from there northward to southern British Columbia, apparently 
always restricted to relatively low elevations. It then recurs in a dis- 
junct area radiating out from the Great Lakes to the James Bay area, 
and to the Gaspé, with a single report from Maine. It is reputed to 
occur eastward to Newfoundland. The explanation of this distribution 
pattern is particularly difficult in view of the fact that F. bolanderi is 
almost exclusively corticolous, with rare (and probably never permanent) 
populations on rocks. It is furthermore "eritical" in that it is clearly a 
non-aretie species, which could not have survived in treeless areas, by 
any stretch of the imagination; it is not known today from any area in 
immediate proximity to the Tundra. As a consequence, the statement 
by Abbe (in Butters & Abbe, 1953, p. 69) is of interest. Abbe suggests 
that the flora of the North Shore of Lake Superior (and specifically that 
of Cook Co., Minnesota) was not replenished, in late-glacial or post- 
glacial time from a ‘‘pool of ‘Cordilleran’ or ‘arctic’ rarities in the Driftless 


226 Rhodora [Vor. 60 


Area." If this statement by Abbe is accepted as true, the existing range 
of Frullania bolanderi becomes incomprehensible. The simplest explana- 
tion for its existing distribution in western North America involves its 
migration into the Lake Superior area from a Pleistocene "refugium" in 
the Driftless Area. The suggestively similar case of Mertensia paniculata 
(Ait.) G. Don? known from the periphery of the Driftless Area (n.e. Iowa), 
the western Great Lakes area, north to James Bay, Quebec; then again 
in the far West from Montana and Idaho to Washington, is equally 
impossible of a simple explanation, if Abbe's hypothesis of preglacial 
migration is uniformly applied. "The Mertensia, like the Frullania, is a 
plant of forested areas, and not of the tundra. As a consequence, the 
suggestion that these species migrated into the Great Lakes area at the 
heels, so to speak, of the retreating glaciers, from a far western “home” is 
difficult to visualize. More probable, in such cases, is the assumption of 
“persistence,” in the classical Fernaldian sense, with the Driftless Area a 
refugium in which these species were able to survive the Pleistocene 
glaciation (while their originally more widespread eastern range was 
otherwise destroyed). 

Frullania bolanderi (and Mertensia paniculata) are significant species 
to consider, from the viewpoint of phytogeography, since they could 
scarcely have survived in the Great Lakes on any hypothetieal nunatak, 
such as has been postulated by Fernald (1935) for the tip of the Keweenaw 
Peninsula. At least, the survival there of these species would appear to 
hinge on the survival of a well-developed forest, a viewpoint which does 
not deserve serious consideration. 

In this connection, the repeated belief by Hultén (1937), in the 
lack of glaciation on “the islands in Lake Superior," or in nunatak 
“districts about the Great Lakes" is also pertinent. Hultén 
(loc. cit., Plate 43) shows such an “isolated” refugium in the 
Nipigon region of Ontario, north of the North Shore of Lake 
Superior. Ashasbeen repeatedly demonstrated by the geologists 
(see review in Butters & Abbe, 1953), no such ice-free refugia can 
be demonstrated. Indeed, the opposite is fairly well-established: 
all of these areas were glaciated during the Pleistocene. As ¢ 
consequence of such evidence, the “persistence” theory, so 
vigorously propounded by Fernald (1925, 1935) has recently 
received almost no support. In any re-evaluation of the ‘per- 
sistence" theory, however, the evidence from such species as 
Frullania bolanderi must also be considered. 

At the time Steere (1937) mapped the range of this species, it 
was known in the east only from obviously glaciated areas.* 

5 The distribution of the various Tracheophyta cited largely derived from the Eighth Edition 


of Gray’s Manual. 
6 Steere (1937) however assigns the eastern stations of the species to “the unglaciated part 


1958] Schuster,— Notes on Nearctic Hepaticae VII 227 


Steere postulated that the distribution of the species was broken 
up into three “remote and isolated ‘islands’: (1) throughout the 
Pacific coast area... (2) around the southwest end of Lake 
Superior, and (3) in the unglaciated part of the Gaspé Peninsula 
of Quebec, with an extension into Maine....This anomalous 
geographical distribution gains still more significance because of 
resemblance to that of many of the vascular plants reported from 
the Keweenaw Peninsula by Fernald.” The distribution of 
these vascular plants, of course, moved Fernald to postulate the 
occurrence of a nunatak area at the head of the Keweenaw 
Peninsula (Fernald, 1935), which was believed to have served as 
a refugium during the late Pleistocene glaciation. Extensive 
collecting by the writer (1946-1950) in the Great Lakes area soon 
showed that F. bolanderi had a much wider range than had been 
suspected. The species occurred westward along the Minnesota 
border to Oak Island, Lake of the Woods; it occurred southward 
to the presumably partially unglaciated southeastern corner of 
Minnesota (Schuster, 1953), and it was also found at many points 
in Wisconsin, Michigan, and into Manitoulin I., Lake Huron, 
Ontario. Lepage (1945, 1953) extended the known range of the 
species northward to the southern corner of James Bay. With 
these extensions, there was a virtual confluence of two of the 
*islands" that Steere had postulated. More significantly, the 
known range of the species was extended southward into the edge 
of the ‘‘Driftless Area," an area about which there is no question 
as to the absence of Pleistocene glaciation. Furthermore, there 
is clear evidence of the existence, in the Driftless Area, of a well- 
developed forest, during the Pleistocene (i.e., from the “boreal” 
rather than arctic nature of the moss flora of the interglacial 
periods). 

These facts prompt the writer to suggest that the eastern 
portion of the range of this species has radiated outward from the 
“Driftless Area." The latter, as a consequence, emerges as a 
possible refugium, postulated as long ago as 1925 by Fernald, 
who at the time was unable to adduce any concrete evidence in 
favor of such a viewpoint. In fact, Fernald (1935) fails to 
mention this earlier hypothesis. 


of the Gaspé Peninsula of Quebec, with an extension into Maine.” Recent evidence suggests 
no such unglaciated area exists. 


228 Rhodora [Vor. 60 


A final matter of interest is the clear demonstration of a con- 
nection between the existing flora of the “Driftless Area," and 
that of the Lake Superior shore. This will be commented on 
again later. 

A second species with a distribution pattern of a nearly similar 
type is Diplophyllum obtusifolium (Hook.) Dumort. (Fig. 13). 
The range of this species approaches that of F. bolanderi in 
several respects: the wide distribution along the Pacific Coast, 
where it is abundant (several stations not drawn in owing to lack 
of knowledge of the exact localities!), with a much more restricted 
range in the Great Lakes area (Minnesota, Michigan, Ontario). 
However, D. obtusifolium differs in its range in several respects 
from F. bolanderi. Unlike the latter, it is considered an oceanic 
species, Persson (1952) describing it as a “‘coastbound species;”’ 
it is also a more widespread species, not endemic to North Amer- 
ica, ranging from western Europe to Iceland, and recently 
reported from Honshu, Japan (Amakawa and Hattori, 1955). 
More significantly, it occurs as a rare disjunct not only in the 
Great Lakes area, but also in the old Appalachian Region where 
it has recently been found. 

The Minnesota, Michigan, Ontario, North Carolina and Ten- 
nessee material, i.e., the eastern population, differs in being pre- 
dominantly or wholly autoecious and in the almost constant 
production of gemmae; in these respects it closely approaches 
D. apiculatum, to which the Minnesota plants were referred by 
Schuster (1953), as var. obtusatum. It seems certain now, how- 
ever, that the var. obtusatum 1s much closer to D. obtuszfolium. 
'The Ontario material was kindly forwarded by Mr. H. Williams; 
it was collected at Cache Lake Outlet in Algonquin Park. The 
Tennessee plants were collected by the author on Myrtle Point, 
Mt. Leconte; the North Carolina plants were collected by the 
author in Linville Gorge, Burke Co. 

The Appalachian populations of D. obtusifolium are unques- 
tionably ancient. Equally ancient is the series of western 
populations, which extend in an are from northern California to 
Attu L, in the Aleutians, southward to Honshu Island, Japan. 
The arc-like, oceanic western range of D. obtusifolium is strikingly 
similar to that of F. bolanderi, which extends from California 
northward to British Columbia, and then reappears on the island 


1958] Schuster,—Notes on Nearctie Hepaticae VI 229 


of Sakhalin (Evans, 1915). Asa consequence, the chief difference 
in the American range of D. obtusifolium, contrasted with that of 
F. bolanderi, lies in the retention of a disjunct Appalachian popu- 
lation in the former, vs. the evident lack of such a population in 
the latter. Also, the rarity of D. obtusifolium in the east makes 
it much more difficult to set up a plausible hypothesis as to the 


Fig.13. Range of Diplophyllum apiculatum (dots) and North American range of Diplophyllum 
obtusifoliums. lat. (the eastern stations representing an undescribed subspecies). The Michigan 
station for D. obtusifolium was reported as D. apiculatum (Steere, 1947); it is based on plants 
collected at Bessemer (M. S. Taylor, 1928). Only two, juvenile, sterile plants have been seen 
with apiculate lobes; these, although hardly certainly determinable, could be referred to D. 
apiculatum. The single fertile plant, however, is not the same taxon; it bears a terminal 
perianth subtended by several cycles of sterile leaves; this perianth-bearing shoot bears innova- 
tions that are androecial (and, at least in one case, paroecious). In other words, the leading, 
gynoecial shoot bears male or bisexual innovations. This feature, together with the uni- 
formly rounded ventral leaf lobes identifies these plants as the eastern race of D. obtusifolium, 


derivation of the Great Lakes population. "The analogies in its 
range to that of F. bolanderi are at least suggestive. 

If Fig. 12 and 13 are compared, a possible hypothesis for the 
origin of the easternmost stations of Frullania bolanderi can be 
derived by analogy to the existing range of Diplophyllum obtusi- 
folium. Is it not possible that these eastern stations of 
F. bolanderi, the so-called “third center” of Steere (loc.cit.), were 
derived from an Appalachian refugium? Although F. bolanderi 


230 Rhodora [ Vor. 60 


is currently unknown from the northern end of the unglaciated 
Appalachian plateau, it would not prove at all surprising to find 
reliet stations for it in this area. 

'The interpretation of the existing range of these two species 
must involve recognition of the fact that both are lowland species 
of forested areas, D. obtusifolium occurring only rarely (in Europe) 
in the higher mountains. Neither species can be interpreted as a 
"nunatak species," however loosely we define this category. 


(4) CORDILLERAN SPECIES EXTENDING EASTWARD AND FOUND AGAIN IN 
THE DRIFTLESS AREA, BUT NOT ALONG LAKE SUPERIOR 


The classical case, previously cited by Steere (1937) is that of Asterella 
saccata (Fig. 14). This thallose, calciphilous species, occurs from the 
Yrkon and Alaska southward to Oregon, Wyoming, New Mexico, and 
probably Mexico, at high elevations. The plant has been found, by the 
Europeans, to be amphizonal in distribution, i.e., with the bulk of its 
range in the arctic-alpine, but a limited and scattered range as an xero- 
thermophyte (see Reimers, 1940). This species, in Europe, shows a 
similar distribution pattern to Atkalamia hyalina (Fig. 15), which is 
treated subsequently. In both cases, there is a small area along the 
Mississippi River, in southeastern Minnesota, where a totally disjunct 
population cecurs. The physica! environment along the bluffs of the 
Mississippi River is so different (basswccd-map!e forest on the rich wooded 
slopes) that it seems difficult to reconcile such distributional anomalies. 

At the present time, no other species of Hepaticae is known to have an 
exactly parallel pattern of distribution. 


(5) ARCTIC SPECIES OCCURRING IN THE DRIFTLESS AREA BUT NOT IN THE 
LAKE SUPERIOR REGION 


During the period 1946-1950, the writer carefully investigated the high 
bluffs on the Minnesota side of the Mississippi River, facing Wisconsin 
to the east. On ledges on these bluffs, usually in rather dense shade, but 
occasionally near the exposed summits of the bluffs, were found two 
species with a wholly anomalous distribution. These occurred here, in 
part, in the shade of basswood-maple forest, on rich mesic slopes, as- 
sociated with such "southern" species as Reboulia hemisphaerica. The 
existing range of these two species may have considerable bearing on 
the problem of the origin of the post-Pleistocene flora of the Great Lakes 
region. 

ATHALAMIA HYALINA (Sommerf.) Hattori (Fig. 15). On a number of 
bluffs facing the Mississippi River, from Waccouta southward, Le., at 
the edge of the “Driftless Area," small to extensive colonies of this arctic- 
alpine species were discovered. ‘The plant is strongly caleiphile. In 
North America, it appears to have a western, Cordilleran “center,” from 
British Columbia and Alberta southward to California and Colorado, at 


1958] Schuster,—Notes on Nearctic Hepaticae VII 231 


| 
! 


M 


Fig. 14. North American range of Asterella saccata. The species is now evidently extinct at 
the Winona, Minn. station (see Schuster, 1953). 


232 Rhodora [ Vor. 60 


Fig. 15. North American range of Athalamia hyalina (= Clevea hyalina). 


high elevations; a disjunct station is reported from the Black Hills area 
of South Dakota (Frye & Clark, 1937, in 1937-47; this report requiring 
verification). In the east, the plant is known from two collections, one 
from the Gaspé, the other from northern Vermont. However, the species 
appears to be much more frequent in the high Arctic, north of the Arctic 
Circle, on both coasts of Greenland, on Ellesmere Island to 82°32’ Lat. N., 
and south to Devon Island.’ 


7 Perhaps a nearly parallel case is that of the essentially arctic-alpine Sedum rosea known 
from s.e. Minnesota in the form of a local race (var. leedyi Rosend. and Moore). S. rosea proba- 
bly includes, fide Fernald (1950) “several so-called but scarcely separable species of w. N. Am." 
This ancient species, however, again occurs as a disjunct in the Appalachian region, from central 
New York to Roan Mt., North Carolina, but is widespread in the eastern North American 
Arctic. from Greenland to the coas: of Maine. Except for the few Appalachian stations, at 


1958] Schuster,—Notes on Nearctic Hepaticae VII 233 


The occasional occurrence of the species in Minnesota with Mannia 
fragrans, a xerothermophyte, at the summits of bluffs (bordering the 
"goat prairies" at the summit and on the western slopes of the bluffs), is 
suggestive of the isolated stations in Europe where the plant occurs as an 
xerothermophyte (Suza, 1938; Reimers, 1940; Müller, 1951-54). It is 
possibly more than coincidental that the only xerothermophyte occurrence 
of Asterella saccata in North America is in the same area. It should be 


5 


ccm op 


Fig. 16. North American range of Mannia pilosa (dots) and the doubtfully distinct M. 
sibirica (crosses). 


noted that the European students who have carefully investigated the 
occurrence of these two arctic-alpine species, also often occurring with 
the "southern" xerothermophytic  Mannia fragrans in Europe, have 
concluded that these represent cases of physiological adaptation by 
Pleistocene relicts. The restriction of the two species to the ‘Driftless 
Area" thus acquires special significance. 


which the plant is a clear relict, S. rosea (L.) Scop., sensu lato, has a distribution pattern very 
closely paralleling that of the Athalamia, although not penetrating to as nearly as high a lati- 
tude as the latter. 

Particularly striking, in this case, is also the connection established between the Driftless 
Area, and the Appalachian Region (in both of which the Sedum occurs as a relict!). 


234 Rhodora [Vor. 60 


MANNIA siBIRICA (K. Müll.) Frye and Clark. (Fig. 16, crosses). This 
species is reputed to be a high arctic type, although it has been collected 
so few times that any conclusion must be regarded as tentative. Virtually 
all of the few collections, from Norway and Siberia, are from the far 
north. The only previous report is of a single collection from Alaska 
(Clark & Frye, 1942). During the years 1947-1950 the writer found this 
species, in typical manifestations, a number of times at the edge of the 
“Driftless Area," and slightly to its north, in the St. Croix R. valley, Le. 
in the same area as Athalamia hyalina, and usually associated with it. 
For exact locality data see Schuster (1953). These reports represent a 
major extension of the species southward.* 

Unfortunately, as has been previously emphasized (Schuster, 1953), it 
is still uncertain whether M. sibirica is fully distinct from M. pilosa the 
latter having a wider but still arctic-alpine distribution (dots on map, 
Fig. 16). If the ranges of the M. sibirica-pilosa complex are combined, a 
nearly continuous series of collections is shown from the Driftless Are: 
north to the Lake Superior shoreline (Fig. 16). 

I have also seen several sterile fragments, mixed with spores, repre- 
senting either this species, or the almost inseparable (when sterile) 
Asterella ludwigii; these plants were collected in lime sinks in Alpena Co., 
Michigan. 


(6) ARCTIC SPECIES OCCURRING SOUTHWARD TO THE LAKE SUPERIOR AREA 
AND IN THE DRIFTLESS AREA 


Although a number of Hepaticae of northern affinity occur in the 
Driftless Area, and again on the Lake Superior shore (Scapania mucronata, 
Preissia quadrata, Tritomaria exsectiformis, ete.), only two taxa with 
an essentially arctic-alpine distribution oecur in both the Driftless Area 
and along the cold shores of Lake Superior. One species is Mannia 
pilosa (Hornem.) Frye and Clark, s. str. As will be noted below, it is 
not clear whether M. sibirica can be kept distinct from this. In either 
eventuality, the same basic pattern remains (Fig. 16). Mannia pilosa is 
known from a station in Greenland, from several in Ellesmere Island, 
and from the Alaska-Yukon region; it occurs south of the Arctic Circle 
oniy in the high Cordilleras along the Alberta-British Columbia boundary, 
in the Gaspé Mountains, and at a single station on. Willoughby Mt. in 
Vermont, It then recurs, locally and usually in only small quantity, in 
western Wisconsin, and in eastern Minnesota and northeastern lowa. 
The latter portion of its range extends from the Driftless Region north- 
ward to the north shore of Lake Superior. This distribution pattern, in 
some ways, serves to unite patterns 1 and 5 into a single unit. The 
implications of this are discussed in the final summation. 

è Dr. H. Crum of the Canadian National Museum has just submitted a specimen of this 
species to me for verification. This was collected at Ft. William, Ontario, near the Lake Su- 


perior shore (May, 1955). 


(To be concluded) 


1958] Rollins,-—New Floras for Parts of N. A. 235 


SoME New FLomas ror Parts or NORTH AMERICA.—Although the 
total number of species of higher plants in the high arctic is not great, 
comparatively, the importance and complexity of the plant cover in 
connection with geomorphie processes is being increasingly recognized. 
This means that an ever larger number of investigators outside of botany 
concerned with research on arctic lands are becoming interested in the 
kinds of plants growing there. The excellent recently published ILLUS- 
TRATED FLORA OF THE CANADIAN ARCHIPELAGO! by A. E. Porsild provides 
a sound coverage of the flora for most purposes. "This work, with neat 
realistic drawings of each species by Mrs. Dagny Tande Lid, provides 
keys to the families, genera and species, brief descriptions and notes 
concerning each species and a map showing the total range in northern 
Canada. 

Another book of outstanding quality concerned with the Canadian 
flora is: FLORA OF MANITOBA? by H. J. Scoggan. Here is a volume that 
ranks with the best state and provincial floras and is superior to most of 
them. The work covers all species and well marked varieties, native or 
introduced, known to occur within the provincial boundaries. There 
are keys to the families, genera, species and varieties and notes concerning 
the habitat and geographical distribution of each species or variety. De- 
scriptions are omitted. The first 37 pages are devoted to the history and 
scope of the flora together with a discussion of the physical features of 
Manitoba, its climate and vegetation and the affinities of the flora. 
Scoggan has critically dealt with literature and specimen records for the 
Province, excluding many names shown to have been incorrectly applied 
or otherwise not authenticated as part of the present-day flora. In an 
analysis of Manitoba's flora, Scoggan finds 1,417 species and 124 sub- 
species or varieties of vascular plants present. This contrasts with 340 
species, subspecies or varieties reported by Porsild for the Canadian 
Archipelago. 


SPRING FLORA OF THE DALLAS-FORT WORTH AREA, TEXAS, by Lloyd H. 
Shinners, covers the ten counties immediately surrounding these two cities, 
but the book will have application to a considerably wider area. Plants 
blooming between January Ist and the first week of June are particularly 
included in the work, but many carly summer blooming types are also 
mentioned. There are keys to the families, genera and species, and notes 
on outstanding characteristics, habitats, geographical distribution and 
blooming dates. Much of the originality in the book is found in the 
appendices where one finds a discussion of technical terms, the use of 
keys, scientific and common names, pronunciation, rules of nomenclature, 
collecting herbarium specimens, natural history of plants, conservation 
and notes on the background of the book. This book is definitely student 


! National Museum of Canada, Bulletin No. 146. pp. 1-209. 1957. Paperbound $2.00. 

? National Museum of Canada, Bulletin No. 140. pp. 1-619. 1957. Buckram $5.00. 

ë Spring Flora of the Dallas-Forth Worth Area, Texas by Lloyd H. Shinners. 514 pp. 1958. 
Paperbound $5.50 ($5.75 by mail. Published by the author and available at S. M.U., Box 473, 
Dallas 5, Texas. 


236 Rhodora [ Vor. 60 


oriented. It is produced by photo-offset printing and the paper covers 
and pages are held together by a plastic spring binding. 

This book, though both seasonally and geographically limited, is of 
considerable importance because it pertains to an area that has not been 
adequately covered by a flora. 

SPRING FLORA OF CENTRAL OKLAHOMA! by George J. Goodman is similar 
in many ways to Shinner's book. It is produced by photo-offset printing, 
has a large page size and the flexible paper cover, together with the pages, 
are held by a plastie spring binding. The book has keys to the families, 
genera and species which bloom before June Ist in the central area of 
Oklahoma and is obviously designed for use by students at the University 
of Oklahoma. Nevertheless, it is a valuable addition to the floras covering 
portions of the United States. There are short notes on habitats and 
geographic ranges under each listed species. As pointed out by Dr. 
Goodman, the area covered is almost entirely in the Permian Red Beds 
and possesses a rich and varied flora. Some 716 species and infraspecific 
taxa are included in the work.—R. C. ROLLINS, GRAY HERBARIUM, 
HARVARD. 


4 Spring Flora of Central Oklahoma by George J. Goodman. Published by the University 
of Oklahoma Duplicating Service, Norman, Oklahoma. 1958. 125 pp. Paperbound $3.35. 


Volume 60, No. 715, including pages 177—208, was issued 6 August, 1958, 


FARLOW REFERENCE LIBRARY 


cc 


Hodova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS | 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR, ( 

IVAN MACKENZIE LAMB 


Vol. 60 September, 1958 No. 717 
CONTENTS: 

Minor Forms of North American Species of Rosa. Walter H. 
CD Pp CSI RN C ARE CUN COD MER TU. Dn 237 


Notes on Nearctic Hepaticae VI. Phytogeographical Relation- 
ships of Critical Species in Minnesota and Adjacent Areas 
of the Great Lakes. R. M. Schuster (Concluded from p. 


FIBER GU Aut getan su. Nis eine ih c: DEES 243 
A New Species of Vicia (Leguminosae) in Florida. R. K. 

SPOR; TOY nd Nobert Kral e.e esena aia renee 256 
Orange-Yellow-Flowered Claytonia virginica. R. C. Rollins .. 258 
Buchloé dactyloides in Illinois. G. Neville Jones ............ 259 
Rorippa sessiliflora in Essex County, Massachusetts. Stuart K. 

ja a E E E a a a 260 


A New Color Form of Solidago sempervirens. Stuart K. Harris 261 
Evolution of Flowering Plants. Gordon P. DeWolf, Jr. (Review) 262 


Grasses New to Illinois and the Chicago Region. John W. 
Thieret and Sidney F. Glassman ................... sss. 264 


The New England Botanical Club, Inc. 


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CARD-INDEX OF NEW GENERA, SPECIES AND 
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For all students of American Plants the Gray Herbarium Card-index 
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tial to scientific libraries and academies and all centers of botanical 
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JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 September, 1958 No. 717 


MINOR FORMS OF NORTH AMERICAN SPECIES 
OF ROSA 


WALTER H. Lewis! 


During the study of the North American rose species east of 
the Rocky Mountains, several unique and distinct forms have 
been observed. Variants include: individuals with more than 
five petals in each flower; those with white rather than pink 
petals; specimens having unarmed floral stems where armature 
is typical; and those with glandular-hispid hypanthia and 
pedicels when the expression for these characters is normally 
eglandular. Since their distinctiveness eliminates any possible 
confusion in naming them, since transplant studies have shown 
that the forms are not expressions of environmental modification, 
and since they may be of considerable importance to other bio- 
logical disciplines (plant breeding, horticulture), it is felt that 
natural variations such as these should be given nomenclatural 
recognition at the rank of form. These several variations are 
doubtless the result of different types of genetic situations such 
as gene mutations, gene combinations controlling expression in 
certain cases (as probably with armature), and others. 

With the new taxa are included new combinations that have 
been modified in view of the evidences obtained from a revision 
of the North American roses (Lewis, 1957a). 


1. R. nitida Willdenow, f. spinosa f. nov. 


R. carolina L., var. setigera Crépin. RuopoRa 2: 113 (1900) pro syn. 
R. carolina x R. nitida Crépin. Rnopona 2: 113 (1900). 

R. nitida x R. palustris Rydberg. N. Am. Fl. 22: 496 (1918). 

? R. nitida x R. virginiana Rydberg. N. Am. Fl. 22: 502 (1918). 


1 The Blandy Experimental Farm, University of Virginia. Present address: Department of 
Biology, Stephen F. Austin State College, Nacogdoches, Texas. 


238 Rhodora [Vor. 60 


Caulis floreus armatus setis tenuibus et amplificatis infrastipulatis spinis 
saepe inparem. Floral stems armed with fine bristles and enlarged 
infrastipular thorns, often in pairs. Very rarely these stems are without 
the bristles typical of R. nitida and have only infrastipular thorns. "The 
kind of armature is the only certain way in which the form may be dif- 
ferentiated from R. nitida. 

HOLOTYPE.— Wilton, Franklin County, Maine. M. L. Fernald 11 
Aug. 1894 (au). 


Both Crépin (1900) and Rydberg (1918) believed f. spinosa to 
be a hybrid between R. palustris Marshall and R. nitida. Con- 
sidering the success of many artificial hybrids between most 
diploid Cinnamomeae species (Erlanson 1934), such a cross 
could conceivably be possible in nature. The F, generation 
might even be male fertile as, for example, has been found in the 
progeny of R. Woodsii x R. blanda and R. blanda x R. pisocarpa 
crosses (Flory 1950). The small amount of abortive pollen 
(16%) found from a specimen of f. spinosa collected at Orono, 
Maine, would add evidence to this opinion. The difficulty 
with this approach arises, however, when it 1s realized that one 
of the parents, R. palustris, is not known from Newfoundland 
or central and northern Nova Scotia (some specimens from the 
latter province have been confused with R. palustris, but they 
are R. virginiana Miller). Since typical specimens of R. nitida, 
f. spinosa have been examined from Newfoundland at St. George 
Bay and at St. John's, it hardly seems possible that the form 
is a result of a cross between R. palustris and R. nitida when the 
variation is found in a region separated by hundreds of miles of 
water from one of the parents. It has been pointed out to the 
author, however, that a bird might transport the hybrid fruit 
from an area where such a hybrid is possible and deposit it in 
an area some distance from one of the parents. Although such 
an event would explain the occurrence of the form in Newfound- 
land, there is no evidence as yet to support this theory. Until 
R. palustris is definitely established as native to Newfoundland, 
experimental hybridization produces a phenotype similar to f. 
spinosa, or until bird migration 1s known to be a significant 
factor here, the form is considered a non-hybrid variation of 
R. nitida. 

Rosa nitida x R. virginiana of Rydberg (1918) tentatively has 
been placed in synonymy under R. nitida, f. spinosa. Erlanson’s 


1958] Lewis,—North American Species of Rosa 239 


(1934) data showed that artificially produced F; plants from 
these species were highly sterile—a condition not apparent in 
the pollen of f. spinosa from Orono, Maine. This plant is 
morphologically similar to those cited by Rydberg as repre- 
sentative of the hybrid, R. nitida x R. virginiana. 


Selected specimens examined:—CaNADA. NEWFOUNDLAND: St. John's, 
Robinson & Schrenk, 3 Aug. 1894! (au). qQuEREC: Stanstead Co.: 
Magog, Pease 728 (au). Unrrep Staves. MAINE: Franklin Co.: Wilton, 
M. L. Fernald, 11 Aug. 1894 (as R. carolina x nitida ? by Crépin; as 
R. nitida x palustris by Rydberg; as R. palustris ? by Fernald) HOLOTYPE 
(au). Kennebec Co: Pine Point, Scarboro, Hyland 321 (MAINE). Penobs- 
cot Co.: Bangor bog, Orono, Fernald 2958 (MAINE). Piscataquis Co.: 
Foxeroft, G. B. Fernald 161 (au), Fernald 319 (au). Washington Co.: 
West Sidney, Hyland 115 (MAINE). NEW HAMPSHIRE: Coós Co.: Pike 
pond, Stark, Pease 23778 (au). 


2. R. palustris Marshall, f. inermis (Regel) stat. nov. 


R. hudsoniana Thory ex Red. Roses 1: 95 (1817). 

R. hudsoniana salicifolia Thory ex Red. Roses 1: 95 (1817). 

R. carolina salicifolia Seringe ex DC. Prod. 2: 605 (1825). 

R. carolina L., var. inermis Regel. Tent. Ros. Mon. 78 (1877). 

R. carolina inermis Schuette. Am. Ass. Adv. Sci. 46: 279 (1898). 

R. palustris Marshall, var. inermis (Schuette) Erlanson. Pap. Mich. Ac. 
Sci. Arts & Let. 5: 90 (1925). 


Individuals typical of R. palustris except that the floral stems are 
without armature. The form occurs infrequently throughout the range 
of the species. 


Selected specimens examined. CANADA. ONTARIO: Muskoka Co.: 
Dorset, C. Huber 5854 (as R. palustris Marsh., var. inermis (Schuette) 
Erl.) (au). Unrrep Srares. connecticut: Windham Co.: Plainfield, 
Wauregan, Sheldon 537 (as R. carolina) (aH). FLORIDA: Putnam Co.: 
Welaka, Laessle, 26 June 1940 (FLAs). MAINE: Hancock Co.: Seal Harbor, 
Mount Desert I., Redfield, 21 Aug. 1890 (as R. carolina) (aH). MASSA- 
CHUSETTs: Berkshire Co.: Lenox, Hoffmann, 11 Aug. 1911 (as R. carolina) 
(an). Norfolk Co.: Milton, Churchill, 17 July 1898 (as R. carolina) (au). 
MICHIGAN: Charlevoix Co.: Boyne City, Erlanson 5789/B (as R. palustris 
var. inermis) (micH). Cheboygan Co.: Lancaster Lake, Ehlers 613 (as 
R. carolina) (aH). NEW vonk: Monroe Co.: Manitou Beach, Killip 7843 
(as R. carolina) (aH). Tompkins Co.: near Spencer Lake, Muenscher & 
Bechtel 520 (as R. carolina) (aH). TENNESSEE: Unicoi Co.: Unaka Springs, 
Lyle 19415 (TENN). VERMONT: Rutland Co.: East Wallingford, Kent, 
June 1897 (as R. carolina) (aH). wisconsin: Brown Co.: Blesh’s Farm, 
Schuette, 22 June 1890 (as R. carolina inermis) (an). 


240 Rhodora [Vor. 60 


3. R. palustris Marshall, f. alba (Rafinesque) stat. nov. 
R. carolina L., var. alba Rafinesque Ann. Gen, Sci. Phys. 5: 214 (1820). 


This form established by Rafinesque (1820) as a variety of 
R. carolina L. (Sp. Pl. 1: 703, 1762) non L. (Ibid: 492, 1753) is 
nomenclaturally corrected in combination with the binomial for 
the eastern swamp rose. No white petaled specimens have been 
observed in this study. 


4. R. palustris Marshall, f. plena f. nov. 


R. carolina L., var. pimpinellifolia Rafinesque. Ann. Gen, Sei. Phys. 5: 

214 (1820) pro parte. 

R. hudsoniana flore multiplici Thory. Prod. Gen. Rosa 147 (1820). 

R. hudsoniana subcorymbosa Thory ex Red. Roses 2: 109 (1821). 

R. carolina L., var. scandens (Thory) Seringe ex DC. Prod. 2: 605 (1825). 

R. carolina L., var. hemisphoerica Seringe ex DC. Prod. 2: 605 (1825). 
A form of R. palustris with more than five petals in each flower. No 

specimens have been observed. 


5. R. Woodsii Lindley, f. hispida f. nov. 

R. adenosepala Wooton & Standley. Contr. U. S. Nat. Herb. 16: 131 
(1913) pro parte. 
R. MacDougali x R. ultramontana Rydberg. N. Am. Fl. 22: 523 (1918). 

Hypanthia sunt glandulari-hispida et plerumque eum pedicellis glandu- 
lari-hispidis. Hypanthia are glandular-hispid and usually with glandular- 
hispid pedicels. Accompanying the glandular-hispid hypanthia and 
pedicels in the type are glandular leaflets, gland-tipped double serrations, 
glandular stipules, and hispid petioles. 

HOLOTYPE: Near Helena, Lewis & Clark County, Montana. F. D. 
Kelsey, June 1892 (as R. Fendleri). Herbarium of Montana State College. 

Selected specimen examined. MONTANA: Missoula Co.: Missoula, 
Sandberg, Aug. 1892 (as R. pisocarpa) (MIN). 


6. R. blanda Aiton, f. carpohispida (Schuette) stat. nov. 


R. blanda carpohispida Schuette. Proc. Am. Ass, Adv. Sci. 46: 279 
(1898). 

R. blanda Aiton, var. hispida Farwell. Pap. Mich. Ac, Sci. Arts & Let. 
2: 25 (1923). 

Rosa blanda which has glandular-hispid hy panthia and pedicels. 
The floral stems may be somewhat bristly. Although this 
deseription approaches that for the individuals intermediate 
between R. blanda and R. palustris, the forma is known to occur 
north of the range of R. palustris on the Gaspé, Quebec 
(Erlanson 1934). 


1958] Lewis,—North American Species of Rosa 241 


Selected specimens examined. MAINE: Penobscot Co.: Greenbush, near 
Cardville, Hyland 816 (as R. blanda, var. hispida) (MAINE). MICHIGAN: 
Houghton Co.: Grand Traverse Bay, Richards 3644 (as R. blanda, var. 
hispida) (MAINE). MINNESOTA: Houston Co.: near Bee, Butters & Rosen- 
dahl 3827 (min). Rice Co.: 10 mi. w. Northfield, Butters & Rosendahl 
2897 (min). Saint Louis Co.: Duluth, Lakela 1482 (as R. suffulta) 
(MiN). Yellow Medicine Co.: Granite Falls, Moyer 358 (as R. humilis) 
(MIN). wisconsin: Ashland Co.: 3. Ashland, Richards 3402 (as R. 
blanda, var. hispida) (MAINE). Door Co.: Little Sturgeon Bay, Schuette, 
17 Aug. 1891 (as R. humilis x R. blanda) (vs). 


7. R. arkansana Porter, f. alba (Rehder) comb. nov. 


R. pratincola Greene, f. alba Rehder. Mitt. Deutsch. Dendr. Ges. 1910 
(19): 252 (1911). 

R. arkansoides Schneider, f. alba Schneider. Ill. Hand. Laubh. 2: 971 
(1912). 

R. heliophila Greene, f. alba Rehder. Mitt. Deutsch. Dendr. Ges. 1915 
(24): 222. 1916. 

R. suffulta Greene, f. alba Rehder. Journ. Arn. Arb. 3: 17. 1921. 


Since the pubescent leafed variation (= R. suffulta Greene) 
of this species is not considered specifically distinct, the white 
petaled form is combined with R. arkansana. 


8. R. arkansana Porter, f. plena f. nov. 


Forma ultra quinque petala in florem. A form typical of R. arkansana 
except that as many as twenty petals are found in a single flower. 


HOLOTYPE. The form is known only from one locality, Woodrow, Sas- 
katchewan and has been distributed by P. H. Wright. Individuals have 
been planted at The Blandy Experimental Farm, Ace. No. 12876-54 
(Lewis 2310). Herbarium specimens are deposited in the U. S. National 
Herbarium (holotype) and the Bailey Hortorium. 


9. R. arkansana Porter, f. setulosa (Cockerell) comb. nov. 


R. pratincola setulosa Cockerell ex Daniels Fl. Bould. Colo., Univ. Mo. 
Stud. Sci. Ser. 11 (2): 148. 1911. 
R. suffulta Greene, f. setulosa Cockerell. Torreya 18: 179 (1918). 


The form has glandular-hispid hypanthia and usually glandular 
pedicels. The isotype was collected by Cockerell at Boulder, 
Boulder County, Colorado. 

Selected specimens examined. coLorapo: Boulder Co.: Boulder, T. D. 
A. Cockerell (as R. suffulta setulosa) 150TvPE (us). El Paso Co.: 5 mi. 
from Colorado Springs, Penfound, 18 July 1924 (coro). Larimer Co.: 


Fort Collins, Harrington 3262 (cono). montana: Big Horn Co.: 6 mi. s. 
Pryor, Charff, 10 June 1953 (monr). 


242 Rhodora [Vor. 60 


10. R. carolina L., f. plena (Marshall) stat. nov. 


R. pennsylvanica plena Marshall. Arb. Am. 136 (1785). 

R. parviflora pleno Ehrhart. Beitr. Nat. 4: 21 (1789). 

R. pennsylvanica var. flore pleno Andrews. Roses 2: No. 102 (1828). 

R. caroliniana var. flore pleno Andrews. | Roses 2: No. 104 (1828). 

R. carolina conglobata 'Trattiniek. Rosae. Mon. 2: 156 (1823). 

R. parviflora (flore multiplici) Thory ex Red. Roses No. 11 (1824). 

R. humilis Marshall, var. plena Best. Journ. Trent. Nat. Hist. Soc. 2: 
1 (1889). 

R. carolina L., var. plena (Marshall) Lynes. Baileya 3: 58 (1955). 


First described in the eighteenth century, the double petaled 
form of R. carolina has recently been nomenclaturally revised by 
Lynes (1955). In this study the rare variation is reduced to 
forma status. 


11. R. acicularis Lindley, subsp. acicularis, f. alba (Nakai) comb. nov. 
R. acicularis Lindley, var. Gmelini (Bunge) Meyer, f. alba Nakai. Bot. 
Mag. Tokyo 30: 241 (1916). 

Among the many specimens of this subspecies examined from Europe 
and Asia, only one had entirely white petals rather than the typical pink. 
The material was collected in Tobolsk, Omsk R. S. F. S. R. (re). Nakai 
(1916) cited the following localities: ''Ham-gyóng austr.: Atok-ryóng 
(Nakai 1825) Cho-tyóng-ryóng (Nakai 1569)." 


12. R. acicularis Lindley, subsp. Sayi (Schweinitz) Lewis, f. plena f. nov. 


Forma ultra quinque petala in florem. A form typical of R. acicularis 
subsp. Sayi (Lewis 1957b) except that as many as fifteen petals occur in 
ach flower. 


HOLOTYPE. The form is known from only one locality, Moose Range, 
Saskatchewan. Herbarium specimens from plants growing at The 
Blandy Experimental Farm (Ace. No. 13386-56) are deposited in the 
U. 8. National Herbarium (holotype) and the Bailey Hortorium. 


ACKNOWLEDGEMENTS 

This paper was completed at The Blandy Experimental Farm, University 
of Virginia, under the guidance of Dr. W. 8. Flory, Jr. whose assistance is 
gratefully acknowledged. To Mrs. Nina Woodford and Mr. Roy E. Shepherd, 
the author is indebted for living plants of several new forms. Throughout 
the paper, translations to the Latin have been done by Miss V. H. Randolph. 

An expression of appreciation is due for the courtesy and cooperation of 
the curators of the following herbaria from which loans of specimens have 
been obtained or otherwise made available: University of Colorado Herbarium 
(cono); Agricultural Experiment Station Herbarium, University of Florida 
(FLAS); Gray Herbarium of Harvard University (GH); Herbarium of the Bo- 
tanical Institute of the Academy of Sciences of the U. S. S. R., Leningrad 
(LE); University of Maine Herbarium (MAINE); University of Michigan 
Herbarium (Mrcu); University of Minnesota Herbarium (mın); Montana State 


1958] Schuster,—Notes on Nearctic Hepaticae VI 243 


College Herbarium (Mont); University of Tennessee Herbarium (TENN); and 
the U. S. National Museum, Smithsonian Institution (US). 


LITERATURE CITED 


Crepin, F. 1900. Note upon a probable hybrid of Rosa carolina L. 
and Rosa nitida Willd. Ruopora 2: 112-113. 

EnRLANSON, E. W. 1934. Experimental data for the revision of the 
North American wild roses. Bot. Gaz. 96: 197-259. 

FrLonv, W. S., JR. 1950. Pollen conditions in some species and hybrids 
of Rosa with a consideration of associated phylogenetic factors. Va. 
Journ, Sei. 1 (N.S.): 11-59. 

Lewis, W. H. 1957a. Revision of the genus Rosa in eastern North 
America: A review. Am. Rose Ann. 42: 116-126. 

—— . 1957b. A monograph of the genus Rosa in North America. 
I. Rosa acicularis. Brittonia (in press). 

Lynes, D. 1955. A double-flowered Rosa carolina. Baileya 3: 58. 

Nakal, T. 1916. Praecursores ad floram sylvaticam Coreanam. VII. 
Bot. Mag. Tokyo 30: 217-242. 

RAFINESQUE, C. S. 1820. Prodrome d'une monographie des rosiers de 
l'Amérique septentionale. Ann. Gen, Sci. Phys. 5: 209-220. 

RYDBERG, P. A. 1918. North American Flora 22: part 6. 


NOTES ON NEARCTIC HEPATICAE VI. PHYTO- 
GEOGRAPHICAL RELATIONSHIPS OF CRITICAL 
SPECIES IN MINNESOTA AND ADJACENT 
AREAS OF THE GREAT LAKES 


R. M. SCHUSTER 
(Concluded from p. 234) 


(T) APPALACHIAN SPECIES OCCURRING NORTHWARD TO THE LAKE SUPERIOR 
REGION AND NEAR OR IN THE DRIFTLESS AREA 


A number of species show this distribution pattern, Diplophyllum 
apiculatum (Fig. 13) and Manna rupestris (Vig. 17) being perhaps typical. 
In both cases the population oecurring near Lake Superior is somewhat 
disjunct, and represents the outlying population. A comparison of the 
distribution of the Appalachian endemic, Diplophyllum apiculatum, with 
that of another species of presumably Appalachian origin, Tsuga cana- 
densis, is not without validity. The occurrence of such species of tem- 
perate occurrence around the shore of Lake Superior (and to the south in 
and near the Driftless Area), strongly suggests that the bulk of the 
distribution of these species is restricted to nonglaciated areas, with the 
restricted range around Lake Superior due to post-Pleistocene migration 
northward from a Pleistocene refugium in the Driftless Area. This 
thesis will be examined in more detail in the summation. 


244 Rhodora [Vor. 60 


DiPLOPHYLLUM APICULATUM (Evs.) Steph. (Fig. 13). This common, 
essentially Appalachian species occurs from southern Maine to the 
southern edge of the Appalachian upland, south to northern Alabama 
and Mississippi (unpublished collections of the writer), and recurs west- 
ward sporadieally to the Ozarks. It has evidently been able to push 
north from the glacial boundary to only a slight degree, and has been 
equally unable to invade the Atlantic Coastal Plain. The range is not 
unlike that of Asplenium cryptolepis (see Fernald, 1935, Map 5). 

An old report from Quebec needs verification and seems unlikely. The 
report of the species from the coast of Hudson Bay (Wynne and Steere, 
1943) is, in the light of our present knowledge, almost certainly erroneous. 

The few stations in the Lake Superior area (one each in Wisconsin and 
Michigan), and that at Taylor's Falls, Minn., just north of the Driftless 
Area, are possibly derived from a population which originated in the 
Driftless Area. It is perhaps not entirely fortuitous that the northwestern 
range of T'suga canadensis closely parallels the range of the Diplophyllum.? 

MANNIA RUPESTRIS (Balb.) Frye and Clark (Fig. 17). "This species, 
unlike the preceding, is strongly restricted to calcareous rocks. It occurs 
in the unglaciated Appalachian Plateau and in the Ozarks, apparently 
in the few places where calcareous rocks occur, and from central New 
York (Schuster, 1949) and southern Ohio south to Tennessee, and to 
Linville Caverns, North Carolina. It is frequent in, and at the periphery 
of, the Driftless Area, in Illinois, Iowa, Wisconsin and Minnesota. ‘Two 
stations occur on the Lake Superior margin in Michigan. Additional 
isolated stations occur in the Gaspé, in Vermont, and in Ontario. How- 
ever, the only points at which the species occurs with any abundance are 
in and near the “Driftless Area," and north of the unglaciated plateau. 

As is quite evident from map (Fig. 17), the post-Pleistocene range of 
this species must be regarded as derived, by limited northeastward 
dispersal, from Pleistocene and probably pre-Pleistocene centers in the 
Driftless Area, and in the unglaciated Appalachian Plateau. The rarity 
in the latter appears chiefly a result of the very localized occurrence of 
suitable, moist and calcareous sites in this region, This distribution 
pattern should be compared with that of Frullania bolanderi (Fig. 12) 
since it gives us a partial model for an explanation of the eastern post- 
Pleistocene range of that species. The same general northeastward, and 
never northwestward, post-Pleistocene ''drift" is observable. This is 
easily understandable on the basis of the mesophytie nature of the 


? The preceding discussion of Diplophyllum apiculatum was written three years before the 
species was actually found in the Driftless Area. In June, 1957, the species was found to be 
abundant on damp, sandstone rocks, associated with Lycopodium selago var. patens, at Rocky 
Arbor Roadside Park, northwest of Wisconsin Dells, Juneau Co. 'To one who had never 
previously collected in the Driftless Area, the consociation of species there was very suggestive: 
the Hepaticae included only species with a wide range in medium to high elevations in the 
Southern Appalachian System, among them Lophozia silvicola, L. incisa, Tritomaria ersecta and 
exsectiformis, Lepidozia reptans, Calypogeia neesiana s. str., Geocalyr graveolens, and such 
mosses as Diphyscium sessile. The entire hepatic flora thus gave the distinct impression of 
species which are widespread at higher elevations in the Blue Ridge system, as well as northward. 


1958] Schuster,—Notes on Nearctic Hepaticae VI 245 


N [S 


Fig. 17. North American range of Mannia rupestris. Within the dotted line the Ozarkian- 
Appalachian region that has not been submerged nor glaciated since the Paleozoic, and the 
Driftless Area. 


i 


Fig. 18. Range of Marsupella paroica (dots), M. sparsifolia (triangles) and Solenostoma 
appalachianum (plus marks). Not all Appalachian stations of the latter drawn in; for its 
distribution see the text. 


246 Rhodora [ Vor. 60 


Hepaticae involved, and of the post-Pleistocene extension of the prairie, 
and prairie habitats, eastward. 


MARSUPELLA PAROICA Schuster (Fig. 18). This very recently dis- 
tinguished species (Schuster, 1957) is locally common in a small area 
from Brasstown Bald, Georgia, northward to Mt. Rogers and Whitetop 
Mt., Virginia (see Schuster & Patterson, 1957). The plant had been 
confused with both M. emarginata and M. sparsifolia (see Schuster, 1953, 
where reasons are given why it cannot go into M. emarginata).  Disjunct 
stations oeeur in Minnesota and in the Huron Mts. of Michigan; the 
former station has been previously reported as M. sparsifolia, an essentially 
arctic-alpine species (triangles on map, Fig. 18); the latter served as the 
basis of the report of M. emarginata from Michigan. 

The Lake Superior stations of M. paroica are hardly explicable except 
on the basis of post-glacial dispersal northward from an unglaciated area. 
Since the species is evidently an old endemic of the unglaciated Appa- 
lachian plateau, the Lake Superior stations must be derived either from 
this latter eenter or from a Pleistocene refugium in the Driftless Area. 
The latter explanation appears more reasonable on the basis of the relative 
distances involved, 

SOLENOSTOMA APPALACHIANUM Schuster (Fig. 18, crosses). This species 
appears to show a very definite affinity to Solenostoma monticola (Hattori) 
comb. n.'? and is perhaps to be regarded as a vicariad of this Japanese 
species. Its relationships are discussed in Schuster (19582). The 
original collections of S. appalachianum were made by the writer in 
1953-54 in two localities: Neddie Creek, a small tributary of the Tucka- 
seegee R., Jackson Co., North Carolina, and near the Whitewater R., 
ast of Joeassee, Oconee Co., South Carolina. In 1957 two further 
collections were made. In June the plant was collected, with capsules, at 
Pictured Rocks, Alger Co., Michigan, just above the shore of Lake 
Superior, and in July-August in the upper reaches of the Whitewater 
Gorge, in Jackson Co., near Beech Gap in Haywood Co., and in Cullasaja 
Gorge, Macon Co., North Carolina; subsequent collections were made in 
the Chattooga R., in both North and South Carolina, and in n.e. Georgia; 
the plant also was found in Big Creek, in n.e. Georgia. It is noteworthy 
that the Appalachian stations all lie in the “coves,” where the old Mixed 
Mesophytie Forest, or derivatives of it occur. 

The distribution of this species closely follows the same pattern as 
that of Marsupella paroica, except for a generally lower altitudinal range. 
It is perhaps significant that this species, like M. paroica, is a pioneer on 
non-calcareous rocks, therefore could have readily survived the Pleistocene 
glaciation on the damp, non-caleareous sandstone ledges and walls of the 
Driftless Area. Habitats very similar to those in which the species 
grows at Pietured Rocks have been found several times in the Wisconsin 
Dells area. 


10 Jungermannia monticola Hattori, Bull. Tokyo Sci. Mus. 11: 33, 1944. 


1958] Schuster,—Notes on Nearctic Hepaticae VI 247 


DISCUSSION AND SUMMARY 


It is always tempting to generalize from distribution patterns 
which show unusual features. By a process of conscious or 
unconscious selection it is also often possible to misplace empha- 
sis, or to derive unwarranted conclusions. Added to this are the 
additional dangers imposed by a fragmentary knowledge of the 
range of the species (the well-known ‘distribution of collectors” 
phenomenon), leading to erroneous conclusions, as, for example 
that of Steere (1937) with respect to some of the so-called ‘‘Cor- 
dilleran" relicts of the Lake Superior regions. In spite of these 
dangers, study of the present ranges of plants remains a fascinat- 
ing, but somewhat hazardous occupation. Since practically all 
of the extant generalizations, valid and invalid, with respect to 
the distribution of temperate and boreal North American plants 
are derived from study of the ranges of the Tracheophytes, the 
present contribution may serve to add significantly to the avail- 
able evidence. As has been pointed out, the small size of the 
Hepaticae, which is surely related to their ability to survive in 
microhabitats, together with their great genetic stability (partly a 
consequence of their high incidence of vegetative reproduction) 
makes them an ideal group from the point of view of phytogeo- 
graphical studies. 

Admitting the hazards involved in generalizing from a limited 
number of selected species, the following tentative conclusions 
appear warranted. (1) There is a distinct, if somewhat tenuous, 
connection between the “old” flora of the unglaciated Appa- 
lachians and that of the Driftless Area (Fig. 13, 17). Perhaps 
through the latter there is an even more tenuous connection with 
the flora of Lake Superior, presumably the consequence of recent 
migration from a Pleistocene refugium in the Driftless Area 
(Figs. 13,18). (2) There isa definite floristic connection between 
the Driftless Area and Lake Superior. Frullania bolanderi (Fig. 
12) and Mannia pilosa (Fig. 16) serve to demonstrate this con- 
nection, and the range of the latter appears to establish a connec- 
tion between the Lake Superior flora and that of the Arctic. 
Parallelisms in the Angiosperms and Pteridophytes are not 
unknown. (3) There appears to be a striking, if tenuous, con- 
nection between the flora of the Driftless Area and the Arctic, 
as demonstrated by Athalamia hyalina (Fig. 15). A close parallel 


248 Rhodora [ Vor. 60 


has been pointed out in the Angiosperms by Rosendahl (1947), in 
Chrysosplenium iowense Rydb., occurring peripheral to the Drift- 
less Area in n.e. Iowa, again in Alberta, and in the high North 
American Arctic. (4) A pronounced connection between the 
Driftless Area, the Great Lakes region, and the far western flora 
is evidenced by the distribution of Frullania bolanderi (Fig. 12). 
The distribution of the herbaceous angiosperm, Mertensia panicu- 
lata, is suggestively similar. 

The preceding cases, each involving the Driftless Area, are as a 
whole suggestive. Of these, the most meaningful perhaps is that 
of the corticolous species, Frullania bolanderi (Fig. 12). In this 
case, at least, the distribution in the east impinges only on one 
area where the species could have survived the Pleistocene glacia- 
tion, in other words, the Driftless Area. Thus there appears to 
be demonstrated a distinet connection between the flora bordering 
the Lake Superior region, and the much warmer Driftless Area. 
Among the Angiosperms, such cases, although rare, are also 
known (Mertensia paniculata, Abies balsamea, Potentilla tridentata, 
Primula mistassinica, Populus balsamea, Thuja occidentalis, ete.). 
Added to this is the fact that there are occasional species, of 
temperate and largely Appalachian, or Appalachian-Ozarkian 
range, which have attained the southern edge of the Lake Superior 
area, and occur in, or near, the Driftless area as well (Figs. 13, 
17, 18). From this the conclusion can hardly be avoided that, 
in some fashion or another, the Driftless Area has served as a 
refugium for some of the plants, otherwise unknown from the 
Midwest, which have in post-Pleistocene times invaded the 
Great Lakes Area. In this connection, one must consider the 
range of the largely Cordilleran Mertensia paniculata, known 
from stations in northeastern Iowa, and southeastern Minnesota; 
the range of Primula mistassinica, known i.e., from a station in 
the unglaciated portion of Illinois; and the range of Potentilla 
tridentata, known i.e., from a station in southeastern Minnesota 
at the periphery of the Driftless Area, in Houston Co. All three 
species are common around Lake Superior. Additional examples 
appear unnecessary." By this the writer would not try to deduce 

n There are also a host of “‘relicts’’ found in the glaciated Mississippi River-St. Croix River 
valley region, running northward from the Driftless Area towards the Lake Superior region. 


Among them are Primula mistassinica (near Stillwater, Minn.), Streptopus roseus var. longipes 
(Fern.) Fassett (Taylors Falls, Wisc.), Dryopteris fragrans var. remotiuscula Komarov (St. 


1958] Schuster,—Notes on Nearctic Hepaticae VI 240 


a wholesale revegetation of the Great Lakes Area, from a hypo- 
thetical refugium in the Driftless region. However, it is proba- 
ble, from the range of such arctic-alpine species as Athalamia 
hyalina, Mannia sibirica and pilosa, and of such Cordilleran 
species as Asterella saccata, that such “rigid” species, of relict 
distribution regionally, were forced this far southward during the 
time of the Pleistocene glaciation. "That they were not, in all 
cases, able to migrate northward recently to the Lake Superior 
region (as, for instance, in the case of Asterella saccata and 
Athalamia hyalina) does not materially alter this fact. "The only 
other explanation, that the ranges of these species in the ''Drift- 
less Area" are the result of chance post-Pleistocene migrations 
would imply too fortuitous a series of coincidences to warrant 
serious consideration. The writer would therefore suggest a 
serious re-appraisal of Fernald's (1925) hypothesis that the 
Driftless Area served as a possible refugium, from which some of 
the disjunct Great Lakes vegetation could have been derived. 
The argument against this by Fassett (1931) based in part on the 
fact that the Cordilleran-Lake Superior Rubus parviflorus does 
not occur in the Driftless Area is not insurmountable, especially 
in view of the fact that another of the Cordilleran-Lake Superior 
species (Mertensia paniculata) does still survive in n.e. Iowa at 
the borders of the Driftless Area. The absence of specific species 
from the “Driftless Area," which are, however, characteristic of 
the Lake Superior region, proves nothing." 

In the case of the larger herbs, or shrubs (such as the Rubus), 
and of trees, which are exposed to the macroenvironment, survival 
in the Driftless Area, especially during the thermal maximum, is 
scarcely to be expected. Smaller herbs (such as Mertensia) or 
even more likely, small cryptogamic plants (such as Hepaticae), 


Croix Falls, Wisc.), Lycopodium selago var. patens (Beauv.) Desv. (Minnesota R. near Mankato 
and near St. Paul, Minn.), Scirpus caespitosus var. callosus Bigel. (Scott Co. and Ramsey Co., 
Minn., in and near the Minnesota R. valley). The occurrence of these plants as relicts in the 
region between the Lake Superior shore (where they are common pioneer or near-pioneer 
plants), and the Driftless Area (where they are now unknown, except for the Primula), suggests 
that they may be the rear-guard of a considerably larger series of species which at one time may 
have migrated northward from the Driftless Area to the shores of Lake Superior. The inverse 
explanation: that they are recent migrants, southward from the Lake Superior shore region, 
appears highly i mprobable. 

12 Deevey (1949, p. 1391) warned specifically against this type of negative reasoning, stating: 
“There is probably no field of scholarly activity where it is so dangerous, and at the same time so 
tempting, to reason from negative evidence, as biogeography.” 


250 Rhodora [ Vou. 60 


which can survive in restricted microenvironments appear more 
able to withstand major climatic changes, and appear to have 
survived in the Driftless Area in larger numbers. This concept 
receives some indirect support from the fact that the Hepaticae 
with arctic and/or Lake Superior affinity and/or Cordilleran 
affinity (Mannia sibirica, M. pilosa, Athalamia hyalina, Asterella 
saccata) are all members of the order Marchantiales, suborder 
Marchantiinae. The Marchantiales, of all the Hepaticae, are the 
only ones with a well-developed xeromorphic structure, adapted 
to long unbroken periods without available water. This is to be 
considered in the light of the fact that of the 175 species of 
Hepaticae found in Minnesota, less than 10% belong to the sub- 
order Marchantiinae. Yet no arctic-alpine members of the Lake 
Superior flora, belonging to the other, less xeromorphic groups, 
occur today in the Driftless Area. Considering some of the above 
distributional anomalies, which have received far from satis- 
factory explanations, the earlier suggestion of Fernald (1925) 
becomes more plausible than his more recent (1935) hypothesis 
that a “nunatak” area at the head of Lake Superior served as a 
refugium during glacial times. 

The above interpretation of the role of the Driftless Area in 
the vegetation of the western Great Lakes Area is also supported 
by other recent sources which have not been drawn on in the 
foregoing discussion. For example, Braun (1950, p. 522-523) 
states that “The development of the Beech-Maple Forest [of the 
“Glaciated North"] was not the result of uninterrupted migra- 
tions from south of the Wisconsin glacial border. "The aspect of 
mixed forest communities of ravine slopes, and the nature of the 
soil of the Driftless Area and adjacent very old drifts to the west 
suggest that this hilly area may have been a Pleistocene refugium 
for an attenuated mixed mesophytie forest which was isolated 
hereabouts in late Tertiary time, or at latest, in pre-Wisconsin 
time. From this refugium, early post-Pleistocene migration 
took place. This is indicated by the early appearance of Fagus 
and Tilia in records of nearby bogs... .”” Miss Braun concludes, 
therefore, that “Thus two migrations, one from the south and 
one from the Driftless Area, met to the north of the Prairie 
Peninsula,” leading to the present distribution of the Beech- 
Maple forest. Implied in this is a much further westward oc- 


1958] Schuster,—Notes on Nearctic Hepaticae VI 251 


currence of Beech during the ‘mid-postglacial humid, warm 
period" with subsequent curtailment of its range during the 
subsequent xerothermic period. During this period, also, a large 
variety of the larger shrubs that may have survived in the 
Driftless Area, such as Rubus parviflorus, may have suffered 
extinction in the Driftless. According to this interpretation of 
Braun, “The Maple-Basswood Forest is...at least in part 
postglacial in origin. It appears to have been derived by climatic 
modification of the late Tertiary or interglacial forest of the Drift- 
less Area. ..." Such an interpretation of postglacial migration 
outward of various vegetational elements has also been used to 
explain the existing range of various other species. Thus 
McLaughlin (1931, p. 286) has suggested that Hypericum 
kalmianum, a slender shrub up to 6 dm. high, endemic to the 
Great Lakes, survived the Pleistocene in the Driftless Area, 
migrating eastward during the early postglacial period along the 
margins of the glacial lakes. A similar explanation could also be 
advanced for the distribution of the very distinetive Great Lakes 
endemic, Cirsium pitcheri, which today is confined to the shores 
of Lakes Michigan, Huron and Superior. 

The above conclusions, with respect to the role played by the 
Driftless Area in the revegetation of the Lake Superior region, 
stand directly opposed to those recently expressed by Abbe (in 
Butters & Abbe, 1953, p. 69). Abbe, indeed, goes so far as to 
consider the presumed migration of species from the Driftless 
Area northward to the North Shore of Lake Superior as requiring 
"far more in the way of botanical legerdemain" than the migra- 
tion patterns he postulates. However, it must be emphasized 
that Abbe dealt with only the Tracheophytes, and largely with 
the Angiosperms. The presence in the Driftless Area of arctic 
and “Cordilleran”’ elements, as regards the Angiosperms, has been 
known for a long time (the case of Rhododendron lapponicum) 
and has been confirmed by recent work [Rosendahl (1947) 
Chrysosplenium | iowense and Sedum rosea; Williams (1937) 
Mertensia paniculata]. This lends credence to the idea (Fernald, 
1925) that the Driftless Area played some part as a source of 
some of the local populations of otherwise arctic and Cordilleran 
species now occurring around Lake Superior, and in the area to 
the north of it. 


252 Rhodor: [Vor. 60 


Abbe (loc.cit., p. 70), reviews the evidence against considering 
the north shore of Lake Superior, or the Keweenaw Peninsula, as 
“refugia.” With this evidence, there is no valid basis for argu- 
ment. However, Abbe also discards the idea of Fernald (1925) 
that some of the relict species (and specifically the **Cordilleran" 
ones) could have survived in the Driftless Area, and migrated in 
the post-glacial period to their disjunct stations in the Great 
Lakes. He states: “It becomes necessary, if a nunatak theory is 
to be maintained as a philosophical necessity, either to relegate 
such refugia to still unexplored areas or to retreat southward to 
the Driftless Area. Either procedure demands that the rarities 
of today migrate across a terrain already heavily mantled by 
vegetation. It is far more reasonable to recognize the limitations 
of these species which restrict them to migration under the more 
favorable conditions for dissemination found in a periglacial 
zone...or along the strand and shore rocks and cliffs of the 
glacial lakes." | Such an explanation, undoubtedly, holds for the 
majority of widespread plants today found around the shores of 
the Great Lakes. However, a question remains as to whether it 
applies to the so-called “rigid” species (sensu Hultén), whose 
ability to compete and migrate actively appears to be exception- 
ally low or to such forest species as Frullania bolanderi. The 
majority of the so-called disjuncts, or "critical" species, are, to a 
greater or lesser extent, rigid species, as was emphasized by 
Fernald. It is in these cases, where it is difficult to visualize 
fluetuating and migrating populations rapidly extending their 
range in an east-west or west-east direction, where the closely 
adjacent Driftless Area may acquire some significance as a 
"refugium." 

The preceding discussion leaves unanswered the question of 
why there is such a marked disjunct element of arctic species 
around the shores of Lake Superior. The first 15 examples cited 
(Figs. 1-10) can be materially supplemented, and represent only a 
portion of the arctic and arctic-alpine species of Bryophyta known 
from the Lake Superior region. The paper by Steere (1937) lists 
a series of equally interesting disjunct arctic mosses and that by 
Thomson (1954) an impressive list of arctic and “high arctic” 
lichens. This arctic flora can be very simply explained as a relict 
or “old” flora, left behind during the immediate post-glacial 


1958] Schuster,—Notes on Nearctic Hepaticae VI 253 


period, in a locally favorable area. The high incidence of arctic 
species around Lake Superior, and the much lower incidence of 
them in the rest of the Great Lakes Region supports this simple 
explanation. Lake Superior, the deepest of the lakes, is also by 
far the coldest, with the slightest winter-summer fluctuation in 
temperature. As a consequence, a narrow, permanently treeless, 
barren border exists, extending inward for only a few score yards 
(and often to a much lesser extent), and up for 10-50 feet in 
elevation above the Lake (rarely much higher), whose existence 
depends on the cooling effect of Lake Superior. This narrow 
strip has been presumed to represent a microclimatically (and, 
because of wave-action, in some cases a microedaphically) con- 
trolled Tundra strip (Schuster, 1953). The vegetation of this 
strip, involving the Hepaticae outlined (species 1-15), together 
with a series not discussed (Lophozia kunzeana, L. hatchert, L. 
groenlandica, L. grandiretis, L. alpestris, Tritomaria quinqueden- 
tata, Solenostoma sphaerocarpum, Tritomaria scitula, Lophozia 
lycopodioides, L. obtusa, and others; for their regional distribution, 
see Schuster, 1953) compares favorably with that of the coastal 
portion of southern and central Greenland, as delineated by 
B^cher (1933) and of coastal Ellesmere Island (Bryhn, 1906- 
1907; Schuster, unpublished). The same rock-pool community, 
formed by Scirpus caespitosus, accompanied by Polygonum 
viviparum, Potentilla tridentata, Pinguicula vulgaris, characterizes 
some of the coastal portions of Greenland." 

It is therefore postulated that the disjunct species around the 
shore of Lake Superior may belong to two types (1) a series of 
"rigid" species, which may have survived the Pleistocene glacia- 
tion in the Driftless Area, and migrated northward the short 
distance involved to the Lake Superior shore, probably in the 
immediate postglacial period, and (2) a series of widespread 
arctic species, not found in the Driftless Area, which represent 
relicts of the immediate post-glacial period, when the Lake 
Superior area was invaded by a tundra-type of vegetation. The 
species of the first class persist in cool, moist microclimates, in 
many cases, in the Driftless Area, in spite of the much warmer and 


13 This affinity is even more strongly suggested by the fact that Lophozia latifolia Schuster 
(see Schuster 1953, 1954) known previously only from this arctic strip along the Lake Superior 
shore in Minnesota, has recently turned up in collections studied f rom Thule, Greenland, as 
well as in material from Swedish Lapland, from Alaska, and from northeastern Ellesmere Island. 


254 Rhodora [ Vou. 60 


drier general environmental conditions currently obtaining there. 

The species of the second class today persist around the Lake 

Superior shore because of the persistence there of localized tundra 

conditions in the vicinity of the spray zone of the lake.—D EPA RT- 

MENT OF BOTANY, UNIVERSITY OF MASSACHUSETTS, AMHERST, 

MASS. 

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1958] Sehuster,—— Notes on Nearctic Hepaticae VI 255 


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Flora of Northern Quebec. "The Bryol. 56: 101-115. 

McLaveuun, W. T. 1931. Preliminary Reports on the Flora of 
Wisconsin, XIV. Hypericaceae. Trans. Wise. Acad. Sci., Arts & 
Letters 26: 281-288. 

MüLLER, K. 1951-1954. Die Lebermoose Europas. Rabenhorst's 
Kryptogamen-Flora 3rd Ed. 6(1): i-xii, 1-756, figs. 1-247. 

Persson, H. 1946. Some Alaskan and Yukon Bryophytes. The 
Bryol. 49: 41-58. 

————— . 1952. Critical and otherwise interesting bryophytes from 
Alaska-Yukon. The Bryol. 55: 1-25. 

Remers, H. 1940. Geographische Verbreitung der Moose im südlichen 
Harzvorland. Hedwigia 79: 175-373, 21 figs. 

ROSENDAHL, C. O. 1947. Studies in Chrysosplenium, with special 
reference to the taxonomic status and distribution of C. iowense. 
RHopona 49: 25-306, pls. 1053-1054. 

SCHUSTER, Rupotr M. 1949. The ecology and distribution of Hepaticae 
in central and western New York. The Amer. Midl. Nat. 42(3): 
513-712, figs. 1-13, pls. 1-18. 

—————. 1951. Notes on Nearctie Hepaticae, II. The Hepaticae 
of the East Coast of Hudson Bay. Bull. 122, Canadian Ntl. Museum: 
1-62, Pls. I-VIII. 

—. 195la. lbid. IV. Scapania spitzbergensis and Scapania 

convexula in North America. The Bryol. 54(3): 162-180, figs. A-B, 

Map 1. 


. 1953. Boreal Hepaticae, A Manual of the Liverworts of 
Minnesota and Adjacent Regions. The Amer. Midl. Nat. 49(2): 
i-v, 257-684, Figs. 1-16, Pls. 1-110. 

. 1954. Notes on Nearctic Hepaticae, VII. Lophozia 
(Dilophozia) latifolia sp. nov. The Bryol. 56: 257-276, pls. I-II. 

1957. Notes on Nearctie Hepaticae, XII. Marsupella 
paroica sp. nov. The Bryol. 60: 145-151. 

. 1957a. Notes on  Nearctie Hepatieae, XV. Herberta. 
Rev. Bryol. et Lichen. 26: 123-145. 

—————. 1958. Boreal Hepaticae, III. Phytogeography. The 
Amer. Midl. Nat. 59: 257-332. 

——————. 1958a. Notes on Nearctic Hepaticae, XV. Two new 
species of Solenostoma with notes on the classification of the genus. 
The Bryol. 61 (in press). 

— AND P. M. PATrERSON. 1957. Noteworthy Hepaticae from 

Virginia. Rnopona 59: 251-259. 


256 Rhodora [Vor. 60 


Srerre, W. C. 1937. Critical Bryophytes from the Keweenaw Pen- 
insula, Michigan. Rnopona 39: 1-14, 33-46, 6 figs. 

Suza, J. von. (1938) 1940. Denkwiirdige Lebermoose des xerothermen 
Gebietes in der Tschecho-Slowakei. Acta Bot. Bohemica 12: 1-68, 
1 pl. 

Tuomson, Joun W. 1954. Lichens of the 1951 foray to the north shore 
of Lake Superior. The Bryol. 57: 278-201. 

WrirrLAKE, E. B. 1954. The Hepaticae of Arkansas. I. The Bryol. 
57: 7-18. 

Witurams, L. O. 1937. A Monograph of the Genus Mertensia. Ann. 
Mo. Bot. Garden 24(1): 1-159. 

Wynne, F. E. anb W. C. Steere. 1943. The Bryophyte Flora of the 
Kast Coast of Hudson Bay. The Bryol. 46: 73-87. 


A NEW SPECIES OF VICIA (LEGUMINOSAE) 
IN FLORIDA! 


R. K. GODFREY AND ROBERT KRAL 


Vicia ocalensis Godfrey & Kral, sp. nov.—Perennis, caulibus ad 12 dm. 
altis, crebris et inter se multum innectentibus, prostratis vel suberectis. 
Caulibus ramisque insignite striatis, sparse pilosis. Foliis 4-6 foliolatis, 
plerumque 6. Foliolis 3-5 em. longis 3-6 mm. latis, subtus sparse 
pilosis supra granularibus, linearibus vel anguste lineari-oblongis aut 
lineari-elliptieis, apicibus rotundo-mucronatis. Racemibus axillaribus, 
longi-peduneulatis, ferentibus ad 18 flores. Floribus 10-12 mm. longis. 
Pediculis calycibusque pilosis, lobis late deltoidis brevi-aeuminatis. 
Petalis albis, quorum cacumina suffundurtur colore pallido-caeruleo ipsa 
lamina plus minusve suffusa omnino colore pallido-caeruleo, | Legumini- 
bus sparse brevi-pilosis aut glabratis, 4-4.5 em. longis 6-8 mm. latis, 
ferentibus ad 8-12 semina. 

SPECIMENS EXAMINED: Florida. Marion Co.: Juniper Creek bank, 
Ocala National Forest, May 18, 1949, Mather m-270 (FrAs); prostrate to 
climbing, succulent-leaved, blue-flowered plants, common on moist sands 
along creek through sandhills, toward east-central boundary of Ocala 
National Forest, April 7, 1957, Kral 4569 (rsu); forming a dense entangled 
mat in roadside ditch bordering hammock, along Juniper Springs Creek, 
northeast of Juniper Springs, May 3, 1957, Godfrey 55537 (rsu—Type). 

Vicia ocalensis is apparently closely related to two other native 
Vicia species of the southeastern United States, V. acutifolia 
Elliott and V. floridana S. Watson. It is readily distinguished 
from both of the latter by its much more robust stature, its 

! Financial support contributing to this research was made available to the senior author by a 
grant from the National Science Foundation (G-2010). Funds for graduate student assistance 


were furnished by the Research Council, Florida State University. These aids are gratefully 
acknowledged. 


1958] Godfrey and Kral,—New Species of Vicia 257 


Fig. 1 and 2 Vicia acutifolia. Fig. 3 and 4 V. ocalensis. Fig. 5 and 6 V. floridana. 
All drawn to the same scale. 


significantly larger leaflets, flowers, fruits, and seeds. Compara- 
tive distinguishing features are given in tabular form below. 
See also Figures 1-6. 


V. acutifolia V. floridana V. ocalensis 

STEMS. Usually not ex- Usually not ex- Mostly up to 12 
ceeding 8 dm. ceeding 6 dm. dm. long. 
long. long. 

LEAFLETS. 2-4, mostly 4, 4—6, mostly 4, 4-6, mostly 6, lin- 
narrowly linear linear-elliptie to ear-oblong or 
to linear, 1.5-3 suborbicular, linear-elliptic, 
em. in length mostly elliptic, 3-5 em. in 
and 1-2 mm. in 1.0-1.5 em. in length and 3-4 
width. length and 3-5 (76) mm. in 

(-8) mm. in width. 
width. 
RACEMES. 2-12, mostly 8- 1-10, mostly 4- 12-18, mostly 15- 


10-flowered. 6-flowered. 18-flowered. 


258 Rhodora [Vor. 60 


V. acutifolia V. floridana V. ocalensis 
FLOWERS. 8-9 mm. long. 5-7 mm. long. 10-12 mm. long. 
LEGUMES. 2.5-3 em. long 1.0-1.5 em. long 4.0-4.5 em. long 
and 5 mm. and 5 mm. and 7-8 mm. 
broad, 8-12- broad, 1-3- broad, 8-12- 
seeded. seeded. seeded. 
SEEDS. 2.0-2.2 mm. in 2.0-2.5 mm. in 3.0-3.5 mm. in 
diameter. diameter. diameter. 


The range of Vicia acutifolia, according to Small (1933), is 
eoastal plain, Florida, Alabama, and Georgia. 

Vicia floridana is restricted to Florida. Specimens in the 
herbaria of the University of Florida and F. S. U. are from the 
following counties: Alachua, Eustis, Hardee, Hillsboro, Lafayette, 
Levy, Marion, Taylor, and Volusia. 

Vicia ocalensis is presently known only from the type locality 
in the Ocala National Forest, Marion County. 

Our field experience and the limited data with herbarium 
specimen indicate that the habitats of the three species are 
closely similar, namely, moist to wet open areas. We have not 
yet encountered more than one species at a given site, however. 

The authors are greatly indebted to Professor Francis R. 
Walton of the F. S. U. Classics Department for his assistance in 
preparing the Latin diagnosis.—DEPT. OF BIOLOGICAL SCIENCES, 
FLORIDA STATE UNIVERSITY. 


ORANGE-YELLOW-FLOWERED CLAYTONIA VIRGINICA.— We are 
so accustomed to seeing the flowers of the Spring Beauty in 
various shades of rose, pink, or sometimes nearly white, that it 
comes as a shock to see plants of such a familiar species with 
orange-yellow flowers and reddish anthers. Yet ten to a dozen 
orange-yellow-flowered plants of Claytonia virginica have been 
found growing together with the usual color form, along the 
Susquehanna Canal in Maryland, by Mrs. George A. Elbert of 
New York City, and she has provided a specimen and photo- 
graphs in color to prove it. The specimen was so carefully 
prepared that it retains the orange-yellow petal color, matching 
the 10 yr 8/10 moderate orange-yellow band on the Nickerson 
Color Fan. The veins of the petals are red, similar in color to 
the anthers. 

In my own experience, I had not seen an orange-yellow-flowered 


1958] Jones,—Buchloé dactyloides in Illinois 259 


Claytonia and I wondered whether this find of Mrs. Elbert’s was 
unique in the genus. A quick perusal of the literature shows 
that orange-yellow-flowered Claytonias are known. Nelson! 
described C. aurea [later? naming it C. flava, because C. aurea 
turned out to be a homonym] from Henry's Lake, Idaho, and 
Greene? named Claytonia chrysantha from Mount Baker in the 
state of Washington. I have not seen specimens of C. flava but 
isotypes of C. chrysantha do not show the yellow color attributed 
to it by Greene. Indeed, St. John‘ points this out and further 
states that there are no real differences between C. chrysantha 
and C. lanceolata Pursh. Nomenclaturally, this yellow-flowered 
plant has been regarded as a species, subspecies’ and form.‘ 
However, not only are there known orange-yellow-flowered types 
in western North America but apparently C. virginica turns up 
with flowers of this color from time to time. In Bartonia,* 
Ball reported Spring Beauties with orange-colored flowers in 
abundance in a meadow near Quakertown, Pennsylvania. It 
wouid be interesting to know more about the geographical range 
of this color form.—R. C. ROLLINS. 


BucHLO# DACTYLOIDES IN ILLINoIs—Buffalo grass, one of the 
most famous and valuable native North American forage grasses, 
was an important constituent of the shortgrass prairies of the 
Great Plains over which vast numbers of bison formerly grazed. 
Its principal area of distribution extends from western Minnesota 
into western Canada and central Montana, and southward 
through eastern Colorado to Arizona, extending into western 
Louisiana, and northward to northwestern Iowa. It has recently 
been found in Peoria County, Illinois. 

This plant is a stoloniferous perennial grass growing in char- 
acteristic colonies often forming a continuous turf of unusual 
toughness, the short stems rising to a height of a few inches and 
bearing curly leaves. In the fall it becomes dry and dull grayish 
green; in both the green and dry condition it is nutritious to 

1 Bull. Torr. Bot. Club 27: 260. 1900. 

? Univ. Wyo. Pub. Bot. 1: 142. 1926. 

? Leafl. West. Bot. 2: 45. 1910. 

4 Res. Stud. State Coll. Wash. 1: 97. 1929. 


5 Ferris in Abrams, Ill. Fl. Pac. States 2: 122. 1944. 
* Bartonia 7:22. 1915. Iam indebted to Dr. John M. Fogg for telling me of this reference. 


260 Rhodora [ Vor. 60 


grazing animals. The thick turf was cut into blocks to build the 
walls of the “sod houses" of early settlers on the western prairies. 

The phytogeographical significance of the discovery of Buchloé 
dactyloides growing spontaneously in Illinois is that it establishes 
the faet that this grass ranged at least as far eastward as west- 
central Illinois, where it is clearly a relict from early post- 
Pleistocene times. Its discovery here by Dr. Chase in 1956 
certainly does not represent a recent extension of range. Al- 
though it is now known to occur in Illinois in only a single locality 
where it occupies a small area in a relatively undisturbed part of a 
cemetery, it is probable that this colony was only one among 
several or many others that persisted in Illinois possibly down to 
the nineteenth century. The fact is nevertheless rather remark- 
able that during one hundred and fifty years of intermittent but 
extensive botanical collection and exploration in Illinois, this 
plant has remained undetected until now. Its recent discovery 
shows again how nearly impossible it is to discover all the species 
of a region even after long study. In this particular instance this 
grass had been completely overlooked, even by the several dis- 
criminating resident students of the flora of central Illinois, in- 
cluding, among others, Frederick Brendel, the distinguished 
author of Flora Peoriana, who studied the botany of the Peoria 
distriet from 1852 to 1912, and Francis Eugene MeDonald, who 
collected extensively in the same area from about 1880 to 1920. 
It remained for the veteran Illinois botanist, Dr. V. H. Chase, 
to discover this species in Illinois at a station about 400 miles 
east of its nearest known occurrence in northwestern Iowa. 

The collection data of Buchloé dactyloides in Illinois are as 
follows: in Springdale cemetery, Peoria, Peoria Co., August 8, 
1956, V. H. Chase 14304, apparently a relic on soil never in cul- 
tivation. Specimens have been deposited in the herbaria of the 
Smithsonian Institution and the University of Illinois.—G. 
NEVILLE JONES, UNIVERSITY OF ILLINOIS. 


RORIPPA SESSILIFLORA IN Essex County, MASSACHUSETTS.— 
When checking over the Cruciferae in the herbarium of the 
Peabody Museum of Salem I found a sheet of an unfamiliar 
species collected in Salem by the Rev. John Lewis Russell in 


1958] Harris,—Rorippa sessiliflora 261 


1859 bearing the notation, *A weed in garden and seen there for 
many years past." The specimen had been given various 
identifications, all obviously wrong. "Taking it to the Gray 
Herbarium I narrowed it down to Rorippa and with the aid of 
Dr. Reed Rollins finally matched it with R. sessiliflora (Nutt.) 
Hitch., a plant of the South and West. While made nearly a 
century ago this still appears to be the only New England 
collection of the species.—STuART K. HARRIS, 

DEPT. OF BIOLOGY, BOSTON UNIVERSITY. 


A NEw CoLoR FORM OF SOLIDAGO SEMPERVIRENS.—Solidago 
sempervirens L., forma citrea Harris, forma nov. Ad formam 
typicam similis, sed floribus pallide citrinis. Like the typical 
form but the florets pale lemon yellow. 

Any botanist finding this conspicuous color form growing with 
normal plants near the parking lot of the Crane Beach Reserva- 
tion in Ipswich, Essex County, Massachusetts, where it is now 
abundant, would have no reason to suspect that it was not native 
to the area. However, this is not the case, the original source 
was in a different county. 

A single plant of the color form was noticed in a colony of 
normal plants growing on open ground in Winthrop, Suffolk 
County, Massachusetts by Mr. Francis Wade in 1942. He dug 
the plant and moved it to his garden in Stoneham, Middlesex 
County, where it prospered and Mr. Wade divided it annually. 
When he moved to Ipswich, he took the clones with him and 
continued to divide them until he now has a considerable number. 
Seedlings coming up resembled the parent plant. Mr. Wade 
then gathered seed in his garden and scattered it on various 
areas about Ipswich where the typical form was growing. Ina 
number of these areas the new form seems to be well established 
and its pale yellow heads contrast sharply with the deep yellow 
heads of the normal plants. More recently Mr. Wade has 
scattered seeds of the color form along the sides of the highway 
between Newburyport and Plum Island but to date none of 
these plants have flowered. 

Mr. Wade brought me material of the new form in September 
of 1954 and I showed it to Dr. Reed Rollins of the Gray Her- 
barium. Suspecting that the plants might be polyploids because 


202 Rhodora [Vor. 60 


of the head size and thickness of the leaves, he had Miss Ann 
Morrill make a cytological study of the material. The pollen 
proved to be normal and the chromosome number was normal 
for the species. 


Material Studied: Massachusetts, Essex County: from clone of original 
plant, garden of Francis Wade, Ipswich, S. K. Harris ,13982 4 October 
1957 (rvrk in GH; isotype in NEBC); garden, Ipswich, Francis Wade 20 
September 1954 (NEBC); near parking lot of Crane Beach Reservation, 
Ipswich, S. K. Harris 13872, 27 September 1957 (NEmc). Stuart K. 
HARRIS, BOSTON UNIVERSITY. 


EvorurtoN or FLOWERING PLANTS.—Aececording to legend, the Ostrich, 
when frightened, buries its head in the sand, apparently reasoning that 
what cannot be seen or heard has no reality. In human affairs this is 
the sort of attitude that continually harks back to the “good old days," 
forgetting the open sewers, the tainted meat, and the little children 
choking with Diphtheria. Needless to say, there should be no tolerance, 
in Science, of Ostrich reasoning. In Biology, the increase of factual 
knowledge, in both the observational and experimental fields, of late 
years, has been truly remarkable. In many cases, however, theoretical 
considerations have not kept pace with the increase in factual data, with 
the unfortunate result that some biologists persist in trying to square the 
modern eorpus of knowledge with theories that were none too happily 
contrived fifty or one hundred years ago. Indeed, perhaps the most 
serious deficiency in contemporary biology is the absence of a sober, 
impartial, eneyclopedic, evaluation and correlation of the facts available 
in the fields of geneties, ecology, comparative morphology, and taxonomy. 

A short time ago there was published a book entitled ‘Features of 
Evolution in the Flowering Plants." The author is Ronald Good, a 
well known phytogeographer, and Professor of Botany at the University 
of Hull (England). On first glance, the book makes an extremely good 
impression, for it is simply and clearly written, and replete with examples 
and tastefully produced illustrations. A second look, however, shows 
that the book is fifty years out of date. The author has chosen to ignore 
most of the modern data of taxonomy, comparative morphology, and 
geneties. In doing so, he has not only denied himself the answers to 
many of his questions (or at least what answers may be available), but 
he has also denied himself the data necessary to frame his questions 
meaningfully. 

As a plant geographer, the author seems to have picked up a nodding 
acquaintance with a great variety of species of flowering plants—and with 
a number of eurrently unpopular hypotheses about the mechanism of 
evolution. Unfortunately, it frequently appears that he does not have 


! Longmans, Green & Co., London and New York, 1956, 30 shillings. 


1958] DeWolf,—Evolution of Flowering Plants 263 


intimate, first-hand acquaintance with the data which he uses. What is 
worse, however, he seems not to have used all of the available sources of 
information. No discussion of the relationships of the monocotyledons 
and dicotyledons, for example, can have much significance if it ignores 
our present knowledge of the vegetative anatomy of the two groups— 
based particularly on the recent work of I. W. Bailey and his collaborators 
at Harvard, V. I. Cheadle at the University of California, and C. R. 
Metealfe at Kew. Neither can any discussion of the interrelationships 
of the higher dicots have much relevancy unless it takes into account the 
work of A. J. Eames and his students at Cornell. Finally, any discussion 
of the evolution of the Compositae or Gramineae should consider the 
genetie studies of Babcock and Stebbins, and their respective collaborators, 
at the University of California. 

Good's main contention seems to be that evolutionary thought is too 
much dominated by the results of statistical analyses of animal popula- 
tions, or by a priori axioms, and not by demonstrable facts. It is true 
that much of our knowledge of the behavior of structural characters in 
populations is derived from the study of laboratory cultures of the fruit 
fly, Drosophila. One of the values of these tiny, bisexual, inseets is 
their extremely short life span, usually completed within two weeks. It is 
perhaps frequently forgotten by students of genetics and evolution that 
Drosophila is biologically comparable only with annual plants which are 
obligately cross-fertilized—types which are generally considered to have 
reached the acme of specialization. They tend to forget that woody 
plants, biennial and perennial herbs, and self-fertile annuals—which 
aecount for the vast majority of flowering plants—may behave very 
differently, both as individuals and as populations. Modern studies 
have shown that, in the north temperate zone, up to 80 per cent of the 
species in a given plant community may have some propensity for vege- 
tative reproduction under natural conditions. Of the 1,500 species of 
plants studied so far, 46 per cent seem to have some faculty for producing 
seed without the need for eross pollination. Finally, of the 15,000 species 
investigated, about 33 per cent seem to be involved in polyploidy. These 
are phenomena unknown in populations of Drosophila. 

Good had an excellent idea, but he didn't carry through. There remains 
a need for an impartial examination of current evolutionary hypotheses. 
There is a fair amount of botanical evidence, experimental and historical, 
which bears on the origin of species and genera. This evidence does not 
seem to support the neodarwinian concept of natural selection. The 
origin of species, or of any other taxonomic category—as distinct from 
the behavior of specific characters—is a matter of history and experiment, 
not speculation. In the literature of botany and horticulture, which has 
been built up over the past two hundred years, there is a wealth of his- 
torical and experimental data about individual plants and populations. 
It is time these data were used.—Gorpon P. DEWorF, JR., CAMBRIDGE 
UNIVERSITY. 


204 Rhodora [Vor. 60 


Grasses NEW TO ILLINOIS AND THE CHICAGO REGION.—Speci- 
mens cited are in the herbarium of the Chicago Natural History 
Museum (F); herbarium of the Illinois State Natural History Sur- 
vey, Urbana (1LLs); herbarium of University of Illinois, Navy Pier, 
Chicago (Np); and United States National Herbarium, Washing- 
ton, D. €. (us). 

Puccinellia distans (L.) Parl. New to Illinois. Many plants of 
this alkali-grass were found in the Proviso railroad yards at 
Northlake, Cook County, Illinois, June 19, 1957, Glassman 4088 
(NP), T'hieret 3534 (F, ILLS, Us). 

Bromus marginatus Nees. New to the Chicago region. This 
brome-grass was collected in a waste place near Hickory Road 
and Kedzie Avenue, Homewood, Cook County, Illinois, June 
17, 1957, Thieret 3503 (v, ILLS). B. marginatus has been collected 
apparently only once before in Illinois. Dr. J. R. Swallen has 
kindly sent us the data for this collection: Fox River valley near 
Elgin, [Kane County], June 30, 1916, Benke 1340 (vs).—Jonwv W. 
THIERET AND SIDNEY F. GLASSMAN, CHICAGO NATURAL HISTORY 
MUSEUM, AND UNIVERSITY OF ILLINOIS, NAVY PIER, CHICAGO. 

Volume 60, No. 716, including pages 209—236, was published 30, September, 
1958. 


pene NON SE 


PE 
N47 


Hovdova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS l 
RALPH CARLETON BEAN 
CARROLL EMORY WOOD, JR, 


( Associate Editors 
IVAN MACKENZIE LAMB 


Vol. 60 October, 1958 No. 718 


CONTENTS: 


Rediscovery of the Genus Neoparrya Mathias (Umbelliferae). 
Witham A. Weber. Sira see ae ee m ULLA 265 


Three New Annual Sunflowers (Helianthus) from the Southwestern 
United States. Charles B. Heier IP.. eeen n aean Tte 272 


Description, Distribution and Ecology of Three Species of Vau- 
cheria Previously Unknown from North America. John L. 
Blum and Robert T.. Walce TEN E R LIE 283 


The New England Botanical Club, Jne. 
8 and 10 West King St., Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


RHODORA.—A monthly journal of botany, devoted primarily to the 
flora of North America. Price, $6.00 per year, net, postpaid, in 
funds payable at par in United States currency in Boston; single 
copies (if available) 60 cents. Back volumes can be supplied at $5.00. 
Somewhat reduced rates for complete sets can be obtained on appli- 
cation to Dr. Hill. 

Notes and short scientific papers, relating directly or indirectly 
to the plants of North America, will be considered for publication. 
Please conform to the style of recent issues of the journal. All man- 
uscripts should be double-spaced throughout. [Illustrations can be 
used only if the cost of engraver's blocks is met through the author 
or his institution. Forms may be closed five weeks in advance of 
publication. Extracted reprints, if ordered in advance, will be fur- 
nished at cost. 


Address manuscripts and proofs to Reed C. Rollins, 
Gray Herbarium, 22 Divinity Ave., Cambridge 38, Mass. 
Subscriptions (making all remittances payable to RHODORA) to 


Dr. A. F. Hill, 8 W. King St., Lancaster, Pa., or, preferably, Botanical 
Museum, Oxford St., Cambridge 38, Mass. 


Entered as second-class matter March 9, 1929, at the post office at 
Lancaster, Pa., under the Act of March 3, 1879. 


INTELLIGENCER PRINTING COMPANY 
Specialists in Scientific and Technical Publications 
EIGHT WEST KING ST., LANCASTER, PA. 


CARD-INDEX OF NEW GENERA, SPECIES AND 
VARIETIES OF AMERICAN PLANTS 


For all students of American Plants the Gray Herbarium Card-index 
of Botanical Names is indispensable. It is a work of reference essen- 
tial to scientific libraries and academies and all centers of botanical 
activity. It includes genera and species from 1885 to date. The sub- 
divisions of species from 1885 to date are now included and from 1753 
to 1886 are in the process of being inserted. Issued quarterly, at $25.50 
per thousand cards. 


GRAY HERBARIUM of Harvard University, 
Cambridge 38, Mass., U. S. A. 


QTRbooora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 60 October, 1958 No. 718 


REDISCOVERY OF THE GENUS NEOPARRYA MATHIAS 
(UMBELLIFERAE) 


WILLIAM A. WEBER! 


Neoparrya Mathias? is a monotypic genus of Umbelliferae 
endemic in the southern Rocky Mountains. N. lithophila was 
described from a single collection of Parry from his 1867 expe- 
dition for the Pacific Railroads (Palmer 1869). Paradoxically, 
the species was never again found in nature until the spring of 
1957, when the writer, having reconstructed Parry’s itinerary 
from his diaries (Parry 1867), succeeded in discovering the actual 
type locality. This locality, given on the original label as “New 
Mexico. On rocks, Huefano Mts, Sept., 1867, C. C. Parry 83", 
is to be found in Colorado, rather than New Mexico as generally 
supposed. 

The morphological affinities of Neoparrya are quite obviously 
with Aletes, a small genus of three species limited also to the 
southern Rocky Mountains. A. humilis is a very restricted 
endemic found at only one station, along Dale Creek in Larimer 
County in north-central Colorado; A. macdougalii occurs on talus 
slopes of the Colorado Plateaus Province in the Four Corners 
region; and the most widespread species, A. acaulis, occupies a 
narrow zone along the eastern base of the Rocky Mountains from 
northern Colorado south into New Mexico, West Texas, and 
northern Mexico. 

Both Aletes and Neoparrya are strongly xeromorphic, with 
thick, glossy, leathery-textured leaves with a basic pinnate form 
but with the leaflets tending to be ternately divided. The leaves 
arise basally from numerous caudices, forming mats of several 


1 University of Colorado Museum, Boulder. 
2 Neoparrya lithophila Mathias in Ann. Mo. Bot. Gard. 16: 393. 1929. 


266 Rhodora Vor. 60 


decimeters diameter. The flowers are yellow in Aletes (in 
Neoparrya they were unknown) and the rays of the umbel in 
both genera tend to be reflexed at maturity. The only feature 
by which Neoparrya seems to differ markedly from Aletes is in the 
oil-tubes which are of various sizes and are scattered through the 
pericarp instead of lying in a uniform row. The ecological 
preference of Aletes is rock ledges and cliffs, usually at altitudes 
of from 6,000 to 8,000 feet, where it forms conspicuous mats on 
the canyon sides. In anthesis the masses of yellow umbels are 
quite showy. 

Aletes acaulis is an abundant and conspicuous species in the 
canyons of the east face of the Front Range in Colorado and New 
Mexico, and Neoparrya is similar enough in its general appear- 
ance, judging from the herbarium material, to indicate similar 
ecological preferences. Nevertheless the plant has eluded col- 
lectors for nearly a hundred years although it must, according 
to Parry’s route, occur in a region rather extensively botanized. 
Dr. Mathias, who described the species in 1929, had hunted it 
unsuccessfully for many seasons in what she felt must be the 
tvpe region, 

Although the writer was unaware of the following quotation 
until after Neoparrya was rediscovered, it is interesting to find 
that Standley (1910) suspected that Parry’s New Mexico local- 
ities of 1867 were incorrectly interpreted, for he says (concerning 
the type locality of Artemisia Parryi Gray): “ ‘Huefano Mts., 
New Mexico, Dr. C. C. Parry in September 1867’. This must 
have been collected in Colorado, for there is a range of this name 
in that state and none, so far as the writer knows, in New Mexico. 
The name should certainly read Huerfano instead of Huefano." 

That Dr. Mathias had independently come to the same con- 
clusion is apparent from her letters to the writer (1956): 

“At the time I published the genus, away back in 1929, I had 
a number of sheets, since the original collection by Dr. Parry 
was a large one and widely distributed; and from the information 
on these sheets and from other notes which I was able to get and 
which I no longer have, I indicated that it was apparently col- 
lected on Huerfano Peak near Servilleta in Taos County. I 
have looked in that area and, so far, without any success. How- 
ever, it is highly possible that it was collected in the canyon of 


1958] Weber,—Rediscovery of Genus Neoparrya Mathias — 267 


the Rio Grande, which is a little inaccessible in that area, as 
vou know. ... I have been on the plateau in New Mexico on 
both sides of the Rio Grande and I have also looked through large 
areas of the eanyon of the Rio Grande and tributary canyons 
where I could get into them. The fruiting specimens were 
collected in September so that I do not believe I have been too 
late. Ihave gone into this area, now, in every month from June 
through late September without success. However, some of the 
perennial Umbelliferae manage to survive without maturing 
fruit over long periods of time and I may have been unfortunate 
in never hitting the area in a good year. It is one of those cur- 
ious plants which I am certain will turn up sooner or later, and 
anything you can do in the way of looking will certainly be ap- 
preciated by us." 

During this long search, Dr. Mathias’ suspicion grew that the 
type locality was incorrectly understood. 

“T had thought originally that it must have been collected 
in the Rio Grande Canyon, not far from Santa Fe or Taos, New 
Mexico. However, last summer when I was at the Missouri 
Botanieal Garden library, I pulled out a number of the old re- 
ports on this expedition and it seems to me that it is just as like- 
ly that this plant may have been collected at the headwaters 
of Huerfano Creek somewhere in the neighborhood of La Veta 
Pass in southern Colorado. If so, it would be on the talus 
slopes, I suspect. I did look over the little Huerfano Peak 
near Walsenburg with a fine-toothed comb last summer, and I 
am certain it is not there. It probably was farther up the head- 
waters of the creek. This is a genus which you might watch for 
when you are in the southern part of the State. I would cer- 
tainly be delighted to find 1t again." 

In a study of the genus /Telianthella (Weber, 1952), the writer 
had a similar experience in settling the type locality of Helianthella 
parryi Gray, and found evidence indicating that any plants which 
Parry collected in the ‘Huefano Mts.’ in 1867, were actually 
collected in Colorado rather than New Mexico, on the head- 
waters of Huerfano Creek. 

Fortunately, several published documents concerning the 1867 
expedition can be used effectively to plot the details of Parry's 
itinerary over the most critical part of the journey. 


268 Rhodora [Vor. 60 


Probably the most important are the Parry notebooks, pre- 
served in the Iowa State College Library, Ames, Iowa. A note- 
book for 1867, pp. 21-39, headed “Route from Fort Wallace to 
Fort Lyon on the Arkansas" records meteorological data taken 
along the course of the journey; dates and place names are linked 
and thus are of value in piecing together the exact route. Most 
of the localities mentioned may be found on the Huerfano Park 
Reconnaissance Map of 1892 (U. S. Geological Survey, 1892). 
The portion of this notebook pertaining to the plant collections 
made in the ‘Huerfano Mountains’ is abstracted below: 

July, 13. Old Fort Lyon; 22. camp 10 mi. above Fort Lyon; 25. Timpas 
Creek near mouth; 26, 27. camp 5 mi. above mouth of Apishapa; 27. 
Apishapa Crossing. ranch 8 miles below mouth of Huerfano; 30. Wilton’s 
Ranch on Huerfano 10 miles above mouth. Doyle's Ranch; 31. Craigs, 
bottom of Huerfano. 

August, 1, 2. bottom of Huerfano; 2, 3. Corral de Toro; 4. camp at 
foot of Sierra Mojada [Wet Mountains]. Greenhorn Ridge, highest point 
reached; 5. two miles above mouth of Apache Creek. Patterson's 
Ranch. on Huerfano bottom; 6. two miles above Butte [presumably the 
Little Huerfano, a prominent volcanic plug in the plain at the mouth of 
the Huerfano]; Badito Ranch. foot of mountains; 7, 8. foot of mountains; 
camp 6 miles above Badito; 9. going to Sangre de Cristo Pass, 9 miles 
below; 10, 11. camp 6 miles above Badito; 11. camp 6 miles east of Pass; 
12. oak grove 10 ft. above ereek; 12-18. Sangre de Cristo Pass, on a bench 
below the erest; 19. survey station, Huerfano Valley; 27-29. Huerfano 
Valley below La Marsea [Mosca] Pass; 31. camp 1 mi. E. of Sangre de 
Cristo Pass; 31-Sept. 3. camp in Sangre de Cristo Valley 2 miles west of 
Pass, 

Sept. 3-4. camp below Placer Creek; 4, 5. camp 3 miles above Fort 
Garland. (The journal ends here). 

The mission of Parry’s group along this route, as will be seen 
from some quotations to follow, was to investigate the feasibil- 
ity of a railroad route through the Sangre de Cristo Pass. For 
this reason a great amount of time was spent in the Huerfano 
Creek drainage, and it was here that the bulk of the Parry plant 
collections were made. The available evidence implies that the 
rest of the journey, from Sangre de Cristo Pass to Fort Garland 
and on to Santa Fe, was a rapid one, which did not permit a 
leisurely study of the vegetation. 

Other valuable tools for interpreting the 1867 collections are the 
narrative by Bell (1870) and the official report on the expedition 
by General Palmer (Palmer 1869). The Bell narrative contains, 


1958]  Weber,—Rediscovery of Genus Neoparrya Mathias 269 


as an appendix, the list of plants collected by Parry, together 
with their general localities (*Colorado', ‘Fort Garland’, ‘Valley 
of the Huerfano’, ‘Sangre de Cristo’ [Pass], ‘Purgatoire Valley’, 
‘Sierra Blanca’, ‘Greenhorn Mountains’, ‘Upper Huerfano’, and 
‘Upper Arkansas’), but Parry implies slight errors, for he claimed 
(Parry 1878) that the list was printed without his having op- 
portunity to revise it. As might be expected, ‘Cynomarathrum 
saxatile Nutt. in herb. Durand’, the specimen subsequently de- 
scribed as Neoparrya lithophila, is listed as having been collected 
near ‘Sangre de Cristo’ [Pass]. 

Despite the fact that Parry’s plant list as published in Bell 
(1870) reports many specimens from Colorado, Porter and Coulter 
(1874) in their Synopsis of the Flora of Colorado, do not cite 
any of Parry’s 1867 collections. Either they did not see these 
specimens or believed them to have been collected, as the labels 
may have indicated, in New Mexico. 

General Palmer’s official report (Palmer, 1869) described the 
‘Huerfano Route’ as follows: 

“This was instrumentally examined by Mr. J. Imbrie Miller, Division 
Engineer, under direction of General Wright. It deflects from the Raton 
Mountain Line at Fort Lyon, and follows up the Valley of the Arkansas 
and its tributary—the Huerfano—to the summit of the ‘Spanish Range’ 
at the Sangre de Christo Pass, 141 miles from Fort Lyon—thence 50 
miles southwestwardly to the Rio Grande, which it intersects near the 
mouth of the Culebro, at a point about 33 miles below Fort Garland, and 
thence down the Rio Grande to Albuquerque. The instrumental exam- 
ination terminated on reaching the Rio Grande, where the elevation was 
found to be 7301 feet above tide—that at the summit of Sangre de Christo 
Pass being 9186 feet. 

The ‘Mosca Pass’ was also surveyed by Mr. Miller, but he reports both 
that and the Sangre de Christo to be impracticable within the Congres- 
sional limit of grade (116 ft. per mile), and there is the additional ob- 
jection of heavy winter snows." 


Bell mentions the Huerfano route, and Parry’s part in the work, 
briefly in the following excerpt (Bell, 1870): 


“Early on the morning of the third day after our arrival in Santa Fe 
[Sept. 16], two of our friends came into the fonda—Calhoun and Imbrey 
Millar,—whom we parted with at Fort Lyon; and before evening all 
Millar's party arrived safe and sound, but much travel-stained and al- 
most shoeless, from their mountain explorations. After leaving us at 
Fort Lyon, they had followed up the Arkansas and its tributary, the 
Huerfano, through the Sangre de Christo Pass to Fort Garland, a mili- 


270 Rhodora [Vou 60 


tary post in the centre of the Rocky Mountains of Colorado; and after 
examining some of the most favorable passes which lead from the heads 
of the Huerfano to the sources of the Rio Grande [Mosca Pass], they 
followed the latter stream for 200 miles down to Santa Fé.” (pp. 148- 
149)... “On the Huerfano Route, Dr. Parry reports that he met with 
no workable coal” (page 114). 


Judging from these several complementary source materials, 
it was abundantly clear that the Huerfano Mountains of Parry 
were not in New Mexico at all, but in Colorado; all that remained, 
then, for a successful solution to the problem of the lost Neo- 
parrya, was a visit to the Sangre de Cristo Pass and the valley 
of the Huerfano, which rises just east of the Pass. 

The actual story of the rediscovery is anticlimactie, for the area 
was accessible, the habitat was indicated, the weather cooperative 
and the season right. Sangre de Cristo Pass was quickly elim- 
inated from consideration, being an overgrazed saddle in the up- 
per montane aspen zone with no promising rock outerops. More 
promising was the eastern approach to the pass along a gravel 
road which follows the base of some huge rock walls and taluses 
in the form of dikes radiating out from a prominent hill south- 
east of Sangre de Cristo Pass. A five-minute reconnaissance on 
the slope of one such dike was sufficient; Neoparrya was there 
in great abundance.? 

As expected, Neoparrya lithophila bears a strong resemblance 
to Aletes acaulis in its caespitose habit and masses of stiff basal 
leaves, yellow flowers and rounded inflorescence with reflexed 
outer rays. Like Aletes, it occurs in crevices of rock outcrops 
with little soil, but despite its north exposure, the slope on which 
the plants grow is more arid than those customarily inhabited by 
Aletes acaulis, rather more like that of A. macdougalii. Asso- 
clates of Neoparrya include Pinus edulis, Leptodactylon pungens 
Hymenoxys richardsonii and Gutierrezia sarothrac. The vellow 
flowers, in late June, were already almost gone, an indication 
of how very late Parry’s September collections were. Many 
mericarps were well developed, and thin sections verified the 
peculiar condition of the scattered oil tubes. Whether these 

3 COLORADO. HUERFANO CO.: forming large clumps up to two feet in diameter among rocks 
on stabilized talus of a dike radiating from Dike Mountain. valley of South Oak Creek, be- 
tween Badito and Sangre de Cristo Pass, 2 miles south and 4 miles west of Badito, ca. 7,000 


ft. alt., 105°05’ W. Long., 37°41’ N. Lat. (Huerfano Park Sheet, Reconnaissance Map, U.S.G.S, 
ed. 1892), 29 June 1957, W. A. Weber & Laurent Gaudreau 10,57 1. 


1958] Weber,—Rediscovery of Genus Neoparrya Mathias — 271 


alone can maintain Neoparrya on the generic level, however, is 
a question which the writer does not feel ready at this time to 
resolve. Suffice it to say that the general morphology, phyto- 
geography, phenology, and ecological preference suggest an ex- 
tremely close relationship with the three species of Aletes. 


REFERENCES 


Bett, W. A. 1870. New tracks in North America, A journey of travel 
and adventure whilst engaged in the survey for a southern railroad 
to the Pacific Ocean during 1867-8. Ed. 2, 565 pp. Chapman & 
Hall, London. 

Ewan, Josepa. 1941. Botanical Explorers of Colorado.—I. Charles 
Christopher Parry. "Trail and Timberline No. 268. 

MaruiaAs, MILDRED E., AND LINCOLN Consrancr. 1944. Umbelliferae, 
in North American Flora 286, part I (Neoparrya, p. 128). 

PALMER, WiLLIAM J. 1869. Report of surveys across the continent in 
1867-68, on the thirty-fifth and thirty-second parallels, for a route 
extending the Kansas Pacific Railway to the Pacific Ocean at San 
Francisco and San Diego. Phila., W. B. Selheimer. 

Parry, C. C. 1867. Journal (manuscript). Iowa State College 
Library, Ames. 

1870. Botany of the region along the route of the Kansas 
Pacific Railway through Kansas, Colorado, New Mexico, Arizona, 
and California. Appendix A, pp. 521-538, in Bell, op. cit. 

. 1878. The Parry botanical collection. Proc. Davenport 
(Iowa) Acad. Nat. Sci. 2: 279-282. 

Porter, T. C., anp Coutrer, Jou M. 1874. Flora of Colorado. 
U. S. Geol. Surv. Miscel. Publication No. 4. 

STANDLEY, PauL C. 1910. The type localities of plants first described 
from New Mexico. Contrib. U. 3. Nat. Herb. 13: 143-227. 

U. S. GEornoarcanL Survey. 1892. Reconnaissance map. Huerfano 
Park sheet. 

WEBER, WILLIAM A. 1952. The genus Helianthella (Compositae). Amer. 
Midl. Nat. 48: 1-35. 


272 Rhodora [Vor. 60 


THREE NEW ANNUAL SUNFLOWERS (HELIANTHUS) 
FROM THE SOUTHWESTERN UNITED STATES 
CHARLES B. HEISER, Jm.! 

I. A NEW AND POSSIBLY EXTINCT SUNFLOWER FROM TEXAS 
Helianthus paradoxus, sp. nov. Herba annua, 15 dm. alta, caule 
sparse scabra, foliis oppositis supra, alternis infra, laminiis inferioribus 
lanceolatis basi euneatis, 10 em. longis, 2.5 em. latis, profunde 3-nervis, 
integris, scabris, petiolis 2 em. longis, pedunculis 3-5, 12-18 em. longis, 
phyllariis 16-20, ovato-lanceolatis vel lanceolatis, sparse hispidis, 10 mm. 
longis, 3 mm. latis, disco ea. 1.8 em. diam., radiis ea. 15, 1.7 em. longis, 
0.7 em. latis, corollis disci purpureis, paleis receptaculi ad apicem glabris. 
TEXAS: Pecos Co., 7 mi. west of Ft. Stockton, Sept. 11, 1947, H. R. Reed 

(type, Us; isotypes, GH, sMU). 

The new species appears to be most closely related to H. 
petiolaris and H. annuus. Its most distinctive feature is the 
strongly 3-nerved lanceolate leaves. The multicellular bases 
of the hairs on the upper surface of the leaf give it a distinct 
punctate appearance. In addition to the leaf shape, the new 
species is readily distinguished from H. annuus by the smaller 
heads, the nearly glabrous stem, and the narrower phyllaries, 
and from H. petiolaris by the glabrous chaff. The tap root is 
poorly developed and its similarity to that of H. agrestis, a 
species of low wet places, suggests that it may be a paludose 
species. 

I have waited several years to describe this species in hopes of 
obtaining additional material. In 1955 an intensive search 
was made for this plant in the area around Ft. Stockton. Seven 
miles west of town presumably where the type was collected, 
there is a low marshy area which I visited in August. Although 
it may have been too early for the species to be in flower, I 
found nothing resembling this species in vegetative condition. 
The area was almost completely covered with the dead stalks 
of another Helianthus, however. A few seeds were secured from 
dried heads and all of them gave rise to plants of H. annuus. 
In 1957 Dr. Raymond Jackson visited this area and also found 
no sunflowers other than H. annuus. 

Two other specimens (Baker 2, GH, TEX) have been seen 
from Fort Stockton which are rather puzzling. These plants 


1 This study was aided by a grant from the National Science Foundation. 


1958] Heiser,— Three New Annual Sunflowers 273 


Fig. 1. Helianthus paradorus. Branch and leaf, x 14; phyllary, x 1!4. Drawing by Ruth 
Smith. 


274 Rhodora [Vor. 60 


are rather similar to H. paradoxus but are taller, much branched 
and have larger heads and leaves (blade 13 em. x 5 em.) Ex- 
amination of the pollen in several flowers revealed the number of 
stainable grains to range from 2 to 37% in contrast to the 98% 
of H. paradoxus. In addition, over one half of the achenes in 
the head were unfilled. It seems likely that these plants are 
hybrids with H. paradorus as one parent and probably H. 
neglectus as the other. The latter is not known to occur in this 
area, however, although it is known from Monahans, Texas, 
a short distance to the northwest. On morphological grounds 
it appears less likely that the second parent could be H. annuus 
which, however, is abundant in the Ft. Stockton area. 

The specimen here designated as H. paradoxus was identified 
by me some years ago as H. practermissus E. E. Wats., chiefly on 
the basis of a specimen which I at that time thought was an 
isotype of H. praetermissus (Heiser, 1952). Subsequent ex- 
amination of the type of this species based on a collection from 
the Sitgreaves Expedition, “head of Rio Laguna" Sept. 27, 
1851, and a collection labeled “Nay Camp, Rio Laguna, N. M., 
August 26, 1851" (Valencia Co.) convinced me that they belong 
to two different species. The latter material appears to be the 
same as H. paradoxus, differing from the type only in slightly 
narrower phyllaries and longer petioles. Helianthus praeter- 
missus differs from H. paradoxus by its very narrow, sessile 
hirsute leaves. The type specimen of H. practermissus is very 
fragmentary and no roots are present so it is not known whether 
it 1s an annual or perennial. 

No specimens of either of these New Mexican sunflowers have 
been secured since the original collection in 1851. It may be 
that both are extinct. Río Laguna, now Río San José, and the 
pueblo of Laguna derived their names from a lake which now 
has disappeared. It is possible that these species were marsh 
plants which grew near the lake. I have made brief excursions 
into this region during two summers and have found only H. 
annuus and H. petiolaris, and Dr. Raymond Jackson of the Uni- 
versity of New Mexico who has made special efforts to collect 
all of the species of Helianthus of New Mexico has so far not 
encountered either H. paradoxus or H. praetermissus. 


1958] Heiser,—Three New Annual Sunflowers 275 
H 


II. DESCRIPTION AND CYTOGENETICS OF A SECOND NEW SUNFLOWER 
FROM TEXAS 

In 1950 Theodore Odell in a search for Helianthus paradoxus 
for me brought back seeds of a sunflower from near Monahans, 
Texas. It was clearly evident that the plants grown from these 
seeds represented a new sunflower obviously closely related to 
H. petiolaris Nutt. In 1955 I was able to visit this area and I 
found these sunflowers growing in scattered colonies along High- 
way 80, ten miles east of Monahans to within two miles of the 
city. 

Helianthus neglectus, sp. nov. Herba annua, 8-15 dm. alta, caule 
dense hispido infra, ramoso supra, foliis inferioribus ovatis, plerumque 
cordatis ad basim, subintegris vel serrulatis remote, sparse hispidis, pilis 
propiter appressis, laminis 7-14 em. longis, 7.5-12.3 em. latis, petiolis 
9-11 mm. longis, pedunculis 10-40 em. longis, phyllariis 25-35, lanceolatis, 
attenuatis 15-24 mm. longis, 2.5-4.0 em. latis, disco ca. 2.3-2.8 em. in 
diam., radiis 21-31, 3.0-3.9 em. longis, 1.0-1.4 em. latis, palei centrale 
receptaculi ad apicem albo-hirsutis. Texas: Ector Co., 10 miles east of 
Monahans, July 22, 1955, Heiser 4123 (type, IND). 

It seems somewhat arbitrary whether this sunflower is treated 
as a species or as a subspecies of H. petiolaris. It does not in- 
tergrade with the latter, but it is closely related as can be seen 
from the densely white pubescent chaff tips in the center of the 
head and most of the other characteristics. It differs from it, 
however, in the very long attenuate phyllaries, the larger heads, 
the broader, and frequently, cordate leaves, the densely hispid- 
hirsute lower stems, and the fact that the branches come off at 
a greater angle. The two species are allopatric, with H. petiolaris 
having a wide distribution over the western half of North Amer- 
ica, and extending within 60 miles of H. neglectus in Martin 
Co., Texas. The new species grows in a more xerophytic area 
than does H. petiolaris in the Texas area. 

The cordate leaf of H. neglectus is similar to that of certain 
races of H. annuus and H. debilis. Helianthus debilis occurs in 
eastern Texas and H. annuus is common throughout Texas, but 
no plants of these two species were seen growing in the area of 
H. neglectus. That H. neglectus owes its origin to past hybridiza- 
tion between H. annuus and H. petiolaris is a possibility, but it 
appears more probable that the new species differentiated from 
the same gene pool as H. petiolaris. The angle of branching in 


276 Rhodora |Vor. 60 


ns = 


N 
N 
IN 
N 
Cm 
S 
\ 
N 
EK 
NN . 


Fra. 2. Helianthus neglectus. Branch and leaf, x 14; phyllary, x 115. 


1958] Heiser,—— Three New Annual Sunflowers 277 
’ 


H. neglectus is unique and is most nearly approached by that of 
H. paradoxus of Pecos County, Texas. 

The new species has been crossed with H. annuus and with 
two or more races of both H. petiolaris and H. debilis. The 
hybrids (Table 1), generally ten plants of each combination ex- 
cept in 5783b where only one hybrid plant was secured, were 
grown in the experimental garden at Bloomington. Pollen 
grains for several plants in each culture were stained in cotton 
blue and those taking a deep stain were counted as “good”. 
Seed set was estimated by examining the number of filled achenes 
in several heads in each plant and in general, the percentage of 
filled achenes was about of the same order as the percentage of 
“good” pollen. All of the species included in the hybridization 
program are diploid (n = 17). Aceto-carmine smears of the 
pollen mother cells from one plant of each of the hybrids were 
made and chromosome pairing was examined at diakenesis. 
Crosses of sister plants of all the species were made and were 
found to give fertile progeny with the expected 17 pairs of chro- 
mosomes. The results of the cytological survey are given in 
Table 1. 

HYBRIDS WITH H. PETIOLARIS.—Helianthus petiolaris com- 
prises two cytogenetic races: race A from the central and western 
United States and race B from the southwestern United States 
(Heiser, unpubl.). Crosses of H. neglectus with both races give 
F, hybrids with varying degrees of sterility and from the exam- 
ination of meiosis it is clear that they both differ from H. neglectus 
by at least one translocation. The hybrids involving race B 
generally show higher pollen fertilities although a few cells in 
these hybrids showed the presence of two chains of four which 
would imply a difference of two translocations. 

The single hybrid secured with H. petiolaris var. canescens, a 
taxon found from southwestern Texas through southern Arizona 
to northern Mexico, shows even greater sterility and indicates 
considerable structural differences in the chromosomes of the 
two parents. 

HYBRIDS WITH H. ANNUUS.— Ten plants of the hybrid, H. 
neglectus x annuus were grown with H. neglectus as the female 
parent. The reciprocal cross failed. Pollen fertility was quite 
low and some heads failed to produce a single filled achene. 


(Vor. 60 


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1958] Heiser,— Three New Annual Sunflowers 279 
Meiosis was examined in one plant but the configurations could 
not be determined with certainty although it was apparent that 
one or more chains were present. These hybrids, in general, are 
rather similar to those of H. annuus x petiolaris. 

HYBRIDS WITH H. DEBILIS.—Helianthus debilis has been shown 
to comprise two cytogenetic races (Heiser, 1956). Representa- 
tives of one group of subspecies, including H. debilis subsp. 
praecox, when crossed with H. neglectus give F; hybrids with 
higher fertilities than does the second group, which includes 
H. debilis subsp. cucumerifolius. Similar results are secured in 
crosses of H. petiolaris with these subspecies. 


III. A NEW SUBSPECIES OF H. PETIOLARIS 


Recent field work combined with garden and herbarium studies 
has convinced me that the southwestern representatives of H. 
petiolaris are quite different from the typical element of the 
species. Crosses of the new subspecies have been made with 
several races of H. petiolaris subsp. petiolaris and it has been 
found that some of the resulting hybrids are fertile whereas others 
show some sterility. It seems desirable to put the subspecies 
on record at the present time, although a detailed discussion of 
the variation and cytogenetics will be deferred until later. 


H. petiolaris subsp. fallax, subsp. nov. Herba annua, 0.5-2.0 m. alta, 
caule hispido vel glabro raro, rubro vel viride, ramoso supra; foliis in- 
ferioribus lanceolatis vel deltoideolaneeolatis, serratis vel integris, 6-15 
em. longis, 1-6 em. latis; pedunculis 5-40 em. longis, phyllariis 12-20 
mm. longis, 3-4 mm. latis, plerumque hispidis, glabris vel hirsutis raro, 
columnis antheris rubris vel purpureis. 

Distribution? western Colorado to Utah south to New Mexico and 
Arizona, 4000 to 8000'; June to Sept. arizona: Apache Co., Painted 
Desert, July 25, 1955. Heiser 4149 (Type, IND); Coconino Co., Eastwood 
and Howell 6925 (vs) ; Cochise Co., Blumer 1488 (Nv); Gila Co., Harrison 
and Kearney 8294 (us); Pima Co., Chondhri 60 (Inv); Navajo Co., Heiser 
4164 (Ind); Yavapai Co., Peebles et al 4291 (vs). cotorapo: Moffat 
Co., Heiser 4540 (1ND); Montrose Co., Walker 152 (au, Us). NEW MEXICO: 
Bernalillo Co., Jackson 2101 (unm); Chaves Co., Griffiths 5685 (us); 
Colfax Co., Heiser 4171a (inp); Dona Ana Co., Dunn 5350 (unm); Eddy 
Co., Whitehouse 16809 (us); Grant Co., Metcalfe 763 (GH, NO, NY, US); 
Quay Co., Fisher 35 (us); Rio Arriba Co., Jackson 744 (UNM); San Juan 
Co., Jackson 2485-2 (unm); San Miguel Co., Standley 5041 (mo, NY); 

2? One specimen from each county is cited. The herbarium abbreviations used are those 


recommended in Index Herbariorum. I would like to express my thanks to the curators of the 
herbaria cited for making the specimens available for study. 


280 Rhodora Vor. 60 


Fia. 3. Helianthus petiolaris subsp. fallar. Branch and leaf, x 14; phyllary, x 11%. 


Heiser,—— Three New Annual Sunflowers 281 


1958] 


Branch and leaf, X 4%; phyllary, X 15. 


Fic 4. Helianthus petiolaris subsp. petiolaris. 


282 Rhodora [Vor. 60 


Sante Fe Co., Heller 3785 (MO, Ny, Us); Socorro Co., Dunn 5090 (UNM); 
Torrance Co., Heiser 4509 (inp); Valencia Co., Weatherwax (IND). UTAH: 
l;mory. Co., Stoutamire 2602 (inp); Garfield Co., Piranian (urc); Grand 
Co., Holmgren and Hansen 3288 (Gu, Ny); Kane Co., Heiser 451.2 (inp); 
San Juan Co., Heiser 4154 (inp); Uintah Co., Rollins 1725 (Gu, MO, NY); 
Utah Co., Jones 509 (Gu); Washington Co., Jones 5257 (mo, us); Wayne 
Co., Stoutamire 2592 (IND). 

Helianthus petiolaris subsp. petiolaris? which is found from 
Wisconsin to British Columbia, south to Texas, and occasionally 
adventive elsewhere, is from 0.5 to 1.4 mm. tall, more branched 
and spreading with stems usually green, rarely red, strigose, or 
somewhat hispid near base, lower leaves usually oblong lanceolate 
to deltoid-lanceolate, usually not over 12 em. long by 6 em. wide, 
peduncles shorter, phyllaries ovate-lanceolate to lanceolate, 
10-14 em. long, 3.5-5.5 mm. broad, strigose, and anthers usually 
purple, rarely red. 

The new subspecies is extremely variable as can be seen from 
the description. From garden cultivation it is apparent that 
some of the variation in nature must be environmental modifi- 
‘ration of the phenotype although a great deal of genetic vari- 
ability is also present. In the eastern part of its range it grades 
into H. petiolaris subsp. petiolaris and in the south into H. 
petiolaris var. canescens A. Gray. In its extreme form it is, 
however, very distinct and if it were not for the presence of in- 
termediates I would consider it a distinct species. The generally 
longer, narrower phyllaries and the hirsute stem are perhaps 
the most reliable characters to separate it from H. petiolaris 
subsp. petiolaris; from H. petiolaris var. canescens it is most 
readily distinguished by a hispid rather than a canescent pu- 
bescence. 

SUMMARY 


Two new species and one new subspecies of Helianthus are de- 
scribed: H. paradorus from Pecos Co., Texas and Valencia Co., 
N. M.; H. neglectus known only from Ector Co., Texas; and H. 
petiolaris subsp. fallar from Colorado, Utah, New Mexico and 


* Nuttall (p. 115, 1821) gives "on the sandy shores of the Arkansa" as the type locality. 
Watson (1929) cites a collection by Nuttall from the “Upper Missouri River" in the Academy 
of Natural Science at Philadelphia as the type. I have examined this specimen and also a 
photograph of a Nuttall specimen from the British Museum labeled ‘‘Arkansa” and the latter 
certainly must be regarded as the type. It is clear from Nuttall's account in his Journal, 
however, that the specimen was collected in what is now northeastern Oklahoma. 


1958] Blum and Wilce,—Three Species of Vaucheria 283 


Arizona. Artificial hybrids of H. neglectus with several other 
species are discussed.—DEPARTMENT OF BOTANY, INDIANA UNI- 
VERSITY, BLOOMINGTON, IND. 


LITERATURE CITED 


Hriser, C. B. 1952. Taxonomic and cytological notes on the annual 
species of Helianthus. Bull. Torrey Club 75: 512-515. 
1956. Biosystematics of Helianthus debilis. Madroño 13: 


145-176. 

NUTTALL, THomas. 1821. A description of some new species of plants, 
recently introduced into the gardens of Philadelphia from the Arkansa 
territory. Jour. Acad. Nat. Sci. Phila. 2: 114-121. 

Watson, E. E. 1929. Contributions to a monograph of the genus 

Helianthus. Pap. Mich. Acad. 9: 305-457. 


DESCRIPTION, DISTRIBUTION AND ECOLOGY OF 
THREE SPECIES OF VAUCHERIA PREVIOUSLY 
UNKNOWN FROM NORTH AMERICA 


Joun L. BLUM AND HoBERT T. WILCE 


During the period from June to November, 1955, a survey 
was made of the distribution and ecology of attached marine 
algae along the coast of the Labrador peninsula. Collections 
were made from the mouth of the Koksoak River, Quebec, north- 
ward along the western side of the Labrador peninsula to Port 
Harvey, Killinek Is., Quebec, and in a less extensive area in the 
regions of Hebron and Saglek, Labrador. Collected materials 
from the mouth of the Koksoak River and False River Bay, 
Quebec, show three species of Vaucheria previously unknown 
from North America.! 

The writers express their gratitude to Dr. I. Mackenzie Lamb, 
Director of the Farlow library and Herbarium, Harvard Uni- 
versity, for the loan of the type material of Vaucheria compacta 
(Collins) Collins, and to Dr. David Irvine, who provided pre- 
served material of V. sphaerospora Nordstedt collected in Scot- 
land. 

Intertidal areas of southeastern and southwestern Ungava 
Bay, especially at the heads of fjords and bays and near the 
mouths of larger rivers and streams, take the form of broad, 


! Specimens of the described species have been deposited in the New York Botanical Garden 
and the herbarium of the University of Michigan. 


284 Rhodora (Vor. €O 


expansive, boulder-strewn mud flats as a result of enormously 
high tides, the low relief of the surrounding watershed, and the 
tremendous discharge of silt from rivers and other streams. In 
such habitats, uncommon in the area investigated, silty mud 
covers much of the shore between tides and, in regions of quiet 
water, extends well into the sublittoral. A dense, wide-spread- 
ing carpet or mat of Vaucheria filaments is common on these 
mud flats. Frequently, colonies of these plants are continuous 
for several square meters or more in extent, penetrating the silt 
and grit only slightly, but becoming strongly infiltrated with it 
so that the mat assumes considerable firmness. The overall ap- 
pearance of the mat is dark green, becoming a glistening black 
when covered with water, and at that time, extremely slippery. 

At the two stations? where these plants were seen the colonies 
were so located that at low water level, in most instances, all of 
the mat would be exposed for varying periods of time. In the 
most sheltered regions, where the water is continually quiet, 
small patches of the mat may extend into the upper sublittoral. 
The latter patches are subject to exposure only during low spring 
tides. 

In addition to the Vaucheria species, a few other marine algae 
are present in this community, but are far less conspicuous. 
Among these are Gloeocystis scopulorum  Hansgirg, Urococcus 
fosliceanus Hansgirg and Calothrix scopulorum Drouet and Daily. 

Vaucheria sphaerospora Nordstedt (fig. 1, 2). This well 
known European species, fruiting abundantly, was collected in 
quantity. It was apparently the dominant element of the mat. 
'The measurements of the Quebec material correspond. well with 
those given by Nordstedt (1878) in his description of this species. 
V. sphaerospora is distinctive in possessing a special fruiting 
branch which bears an oogonium and a single antheridium which 
curves toward the oogonium. This species is the only Pilo- 
boloidean Vaucheria possessing this combination of characters. 

The oogonium is formed from the fruiting branch immediately 
below the suffultory cell which subtends the antheridium, much 
as in V. intermedia Nordstedt and V. minuta Blum and Conover. 
As the cylindrical oogonial filament becomes laterally distended 


2 Wilce 830, September 1, 1955, near the mouth of the Koksoak River, eastern shore; Wilce 
544, September 4, 1955, toward the head of False River Bay on the western side. 


1958] Blum and Wilee,— Three Species of Vaucheria 285 


Fig. 1-17 Vaucueria. Fig. 1 & 2 Vaucheria sphaerospora. Fig. 3-8 V. compacta; fig. 3-5 
drawn from type material in the Farlow Herbarium of Harvard University; Mystic River 
marshes, Malden, Mass., Sept. 1897 (F. S. Collins 477); fig. 3 Antheridium; fig. 4 & 5 Oogonia; 
fig. 6-8 drawn from material collected in Great Pond, Falmouth, Mass. (J. T. Conover, 53-6) ; 
fig. 6 & 7 Oogonia; fig. 8 Antheridium. Fig. 9-12 V. compacta var. koksoakensis; fig. 9 & 10 
Antheridia; fig. 11-12 Oogonia. Fig. 13-17 V. submarina; fig. 13-15 Oogonia; fig. 16 & 17 
Antheridia. All figures about 90X. 


286 Rhodora Vor. 60 


in the formation of the oogonium, the antheridium and the sub- 
tending suffultory cell lose their terminal position and thus ap- 
pear to constitute a lateral offshoot of the mature oogonium 
(fig. 2). A wide terminal pore eventually forms at the upper end 
of the oogonium. The spherical oospore possesses at maturity 
a wall varying from 1—4u in thickness and does not quite fill the 
oogonium. The antheridium is fusiform or conical at maturity, 
its upper end narrowing to its terminal pore. Other pores, num- 
bering 1-4, are at the ends of the papillae, one or more of which 
may equal or exceed the terminal papilla or upper end of the 
antheridium in length. The entire antheridium lies parallel to 
and closely adjacent or appressed to the oogonium. The ter- 
minal papilla of the antheridium curves toward the upper end 
of the oogonium and the papillae other than the terminal one are 
mostly found on the side of the antheridium adjacent to the 
oogonial pore, and are more or less directed toward it. 

Vegetative filaments 28-534 in diameter; oogonium 87-178 x 214-3864; 
oospore 86-1444 in diameter; antheridium 38-57 x 128-1574; antheridial 
papillae 14-33 (-45)u in length; antheridial suffultory cell 50-574 in 
length. 

Vaucheria compacta var. koksoakensis var. nov. (fig. 9-12). 
A typo differt in oogonia longiora. Differs from the type in the 
greater length of the oogonia. 

V. compacta (Collins) Collins has been colleeted in eastern 
United States (Collins 1900, Blum and Conover 1953), and in 
western Europe (vide Christensen 1952). It is dioecious, with 
both oogonia and antheridia borne at the ends of short erect 
branches which occur in short series arising from the vegetative 
filaments. The oogonium is essentially cylindric but is greatly 
swollen at its upper end where the spherical oospore is present. 
Below the oospore the mature oogonium is empty. The stalk 
which bears the antheridium occasionally sends out a branch, in 
such a way that the antheridial branch bears two antheridia 
(fig. 10).  Antheridia dehisce by two or three pores which ter- 
minate conical papillae, one of which is always terminal. 

In the type material of V. compacta (fig. 3-5), and in the mate- 
rial of V. compacta from Massachusetts (J. T. Conover 53- 6, 
east shore marsh, Great Pond, Falmouth, Barnstable Co., Jan. 
31, 1953, fig. 6-8), the length of the oogonium averages 1.5-2.5 


1958] Blum and Wilce,— Three Species of Vaucheria 287 


Xx the length of the oospore (fig. 4-7). In the material of V. 
compacta var. koksoakensis the oogonium is much longer, averag- 
ing from 3-4 X the length of the oospore (fig. 11-12); thus its 
appearance is significantly different from that of the type. Al- 
though this difference could be due to environmental factors, 
on the basis of the available data it seems nevertheless preferable 
not to regard this material as typical V. compacta. 

Measurements of the Quebee material are as follows: Vegetative fila- 
ments 21—50u in diameter; oogonium 87-128 x 235-3574; oospore 92-100 
(-114)u; antheridium 24-48 x 128-1864; papilla of antheridium 8-274 
in length; suffultory cell 28-654 in length; total length of the oogonium 
with its subtending stalk 214-4294. 

Vaucheria submarina Berk. sensu De Wildeman 1899 (fig. 
13-17). Adequate knowledge of the widespread V. dichotoma. 
Ag. and of its forms or varieties, if any, awaits further study. V. 
submarina is certainly closely related to V. dichotoma and may be 
referable to one of the described forms of the latter, such as 
Hauck's V. dichotoma f. marina, although the brief original de- 
seription of the latter form renders the name essentially am- 
biguous. Berkeley, in an even less satisfactory description 
than Hauck’s, raised V. dichotoma f. marina to specific rank, 
V. submarina Berk. De Wildeman, in his Algues de la Flore de 
Buitenzorg (1900) used the name V. submarina Berk. in a precise 
way, and from De Wildeman's description it is possible to iden- 
tify V. submarina as one of the species from the Quebec collec- 
tions. These plants seem to correspond in all essential respects 
with the material collected by De Wildeman in Java. Our 
material was found as a sparse admixture in the V. sphaerospora 
stratum. 

This species, like V. dichotoma, is apparently dioecious, the 
subspherical oogonia being borne singly (in our material) on a 
short stalk directly upon the horizontal vegetative filaments, 
and the bottle-shaped, fusiform or eylindric antheridia being 
borne in uni- or multilateral series, on filaments separate from 
those which bear oogonia. Our material is strikingly smaller 
than V. dichotoma in the size of the vegetative filaments, and is 
further unlike that species in the spherical shape of the oospore, 
which fills a relatively smaller portion of the oogonium. In V. 
dichotoma the entire oogonium is usually filled by the oospore. 


288 Rhodora [Vor. 60 


Vegetative filaments (35-) 50-84 (-100)u in diameter; oogonium 
185-228 X 186-3144; oogonial pore about 12-40u in diameter; oospore 
(130-) 171-186 x (150-) 171-2004; antheridia 43-71 x 114-2004. 


SUMMARY 


Collections of tidal Vaucheriae gathered in the Ungava Bay 
region of northern Quebec are described and brief notes on their 
habitat are given. Vaucheria sphaerospora and V. submarina are 
considered to be new to the North American flora. A collec- 
tion of V. compacta characterized by unusually long oogonia is 
described as V. compacta var. koksoakensis.—CANISIUS COLLEGE, 
BUFFALO, N. Y. AND DEPARTMENT OF BOTANY, UNIVERSITY OF 
MICHIGAN, ANN ARBOR, MICHIGAN. 


LITERATURE CITED 

Buu, J. L. AND J. T. Conover. 1953. New or noteworthy Vaucheriae 
from New England salt marshes. Biol. Bull. 105(3): 395-401. 

CHRISTENSEN, T. 1952. Studies on the genus Vaucheria. I. A list 
of finds from Denmark and England with notes on some submarine 
species. Bot. Tidskr. 49: 171-188. 

Couns, F. S. 1900. Notes on algae. II. Ruopona 2: 11-14. 

Norpstept, O. 1878. Algologiska smasaker. I. Bot. Notiser 1878: 
176-180. PI. 2. 

De Witpeman, E. 1900. Les algues de la flore de Buitenzorg. xi, 457 
pp., 16 pl. Leide. 


Volume 60, No. 717, including pages 237-264, was published 2 October, 
1958. 


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JOURNAL OF THE 


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IVAN MACKENZIE LAMB 


Vol. 60 November, 1958 No. 719 
CONTENTS: 
Striga asiatica and Chrysoma pauciflosculosa in the Carolinas. 
b vun D. LOTU eo E dd Mes 289 
Fleld and Herbarium Studies in Southern Illinois. 
Robert H-"Mohlenbrockz dt. ovo os ath ke ce 292 
Twelfth Report of the Committee on Plant Distribution. 
R. C. Bean, A. F. Hill and R. J. Eaton........... lese. 297 
Notes on the Distinction between the Broad- and Narrow-leaved 
Antennarias of Minnesota. Thomas Morley................ 306 
Euphrasia micrantha New to North America. F.J. Hermann... 307 


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STRIGA ASIATICA AND CHRYSOMA 
PAUCIFLOSCULOSA IN THE CAROLINAS 


Epwarp E. TERRELL 


Striga asiatica (L.) Kuntze, Witchweed, (Scrophulariaceae), 
is parasitic on the roots of corn, sorghum, sugar cane, and other 
grasses. It has long been a serious pest in tropical and sub- 
tropical areas of the Old World. 

Garriss and Wells (Plant Disease Reporter 40(10):837—839. 
1956) reported the occurrence of a species of Striga in Robeson 
County, southeastern North Carolina. Subsequently identified 
by Dr. S. F. Blake as S. asiatica, this was the first record of the 
species in the Western Hemisphere. Since this report intensive 
research was initiated by the Department of Agriculture and by 
agriculturalists and plant pathologists in North Carolina and 
South Carolina to find means of eradicating this pernicious 
weed. Studies of effects of soil types and soll temperatures on 
growth were made (Nelson, Plant Disease Reporter 42(1): 152— 
155. 1958). The same author (loc. cit. 42(3):376-382. 1958) 
stated that 45 species of cultivated and wild grasses may serve as 
hosts for Striga. General information, much of which came from 
earlier studies by South African researchers, was summarized 
by Nelson (loc. cit. 41(5):377-383. 1957), and in U. S. Depart- 
ment of Agriculture, Agricultural Research Service, Special 
Report 22-41, 17 pages, June 1957. The latter publication in- 
cluded drawings of Striga. Seeds of Witchweed may remain 
viable in the soil for twenty years; a single plant may produce 
50,000 to 500,000 seeds, each about 0.2 mm. in length. Ob- 
viously, such seeds are readily disseminated by wind, and once 
present in the soil constitute a long-lasting source of new plants. 


290 Rhodora [Vor. 60 


Witchweed causes stunting and withering of its host; if heavily 
infested, a field of corn may be a complete economic loss. 

At present Striga is known only in the two Carolinas; it seems 
quite possible that it may spread in these two states and possibly 
to adjoining states, despite the efforts to eradicate it. Although 
nothing is known of the time and place of introduction, uncon- 
firmed statements by farmers living in the area of infestation sug- 
gest that the species may have been present there since 1950, 
or even as early as 1946. As cited below, the present author 
collected the species in Robeson County in July, 1955, actually, 
as it turned out, antedating the collection reported by Garriss 
and Wells by about one year; therefore, this was the first collec- 
tion in the Western Hemisphere. Immediately following the re- 
port by Garriss and Wells, surveys by practical botanists in the 
later summer and autumn of 1956 determined that Witchweed 
was then present in at least four counties in southeastern North 
Carolina and in four adjoining counties in northeastern South 
Carolina. This infestation was almost entirely in corn fields. A 
survey by the Plant Pest Control Division, U.S. D. A., revealed 
that Striga was by the end of 1957 present on 1277 farms in eleven 
counties of North Carolina, and on 416 farms in six counties of 
South Carolina (H. R. Garriss, letter dated April 29, 1958). 
These counties were Bladen, Columbus, Cumberland, Duplin, 
Harnett, Hoke, Pender, Richmond, Robeson, Sampson, and Scot- 
land in North Carolina; in South Carolina Darlington, Dillon, 
Florence, Horry, Marion, and Marlboro. 

The present author's collections of Striga asiatica may be 
cited as follows: dry, sandy soil at border of corn field, 3 miles 
east southeast of Pembroke, in Moss Neck community, Robeson 
County, North Carolina, July 28, 1955, E. E. and Bessie Z. 
Terrell 2973(a). A subsequent collection, 2973(b), of the same 
population in fruiting condition was made on August 25, 1955. 
Duplicates of these collections are in the herbaria of the Uni- 
versity of North Carolina (Ncv) and Gray Herbarium (GH). 

Chrysoma pauciflosculosa (Michx.) Greene is a shrubby com- 
posite closely related to or possibly congeneric with Solidago. 
It is here reported for the first time from North Carolina. Chap- 
man (Fl. S. U. S., ed. 2, 1883) listed it under the name, Solidago 
pauciflosculosa Michx., as ranging from ‘Florida to South 


1958] Terrell,—Striga asiatica and Chrysoma pauciflosculosa 291 


Carolina". Small’s Manual (1933) stated, ‘Coastal Plain, 
Fla. to Miss. and S. C." Despite these two listings, no speci- 
mens from South Carolina have been located; therefore, so far 
as known, the collections of Freeman and Radford, cited below, 
represent the first from South Carolina, also. 

At its only North Carolina station Chrysoma grows in deep 
sand under and among turkey oak (Quercus laevis Walt.). It 
is very abundant over an area of several acres and is, in fact, 
the dominant shrub cover. "Typical sand hill species associated 
with it include Selaginella arenicola Underw. subsp. acanthonota 
(Underw.) Tryon, Aristida stricta Michx., Cnidoscolus stimulosus 
(Michx.) Gray, and Euphorbia Ipecacuanhae L. Other sand 
hills near this one were visited, but Chrysoma was absent. Col- 
lections from North Carolina were as follows: dry sand hills, 
114 miles west of Pembroke, Robeson County, June 4, 1955, 
E. E. Terrell 2953(a), (Ncsc); Sept. 4, 1955, 2953(b), (Ncsc, 
Neu, GH); July 17, 1956, 3109, (ncu). The two collections 
(herb. xcv) from South Carolina were: near junction of state 
highways 6 and 602, turkey oak barrens, Lexington County, 
Oct. 10, 1956, O. M. Freeman 56833; sandhill, 4 miles northwest 
of Edmund, Lexington County, Oct. 5, 1957, A. E. Radford 
29849. There is every indication that these two collections 
came from the same locality, which is in central South Carolina, 
about 125 miles southwest of the station in southeastern North 
Carolina. 

It may be noted here that the usual flowering time for this 
species is in September and October. The writer's collections, 
2953(a) and 3109, were made at times when a few plants were 
flowering much ahead of the remainder of the population. 

Appreciation is hereby expressed to the following persons for 
data on distribution and for general assistance: Dr. R. L. Wilbur, 
Duke University; Mr. H. E. Ahles and Dr. A. E. Radford, 
University of North Carolina; Mr. O. M. Freeman, Tryon, 
North Carolina; Dr. H. R. Garriss, Plant Pathology Extension, 
North Carolina State College.— GUILFORD COLLEGE, NORTH 
CAROLINA. 


292 Rhodora [Vor. 60 


FIELD AND HERBARIUM STUDIES IN 
SOUTHERN ILLINOIS 


HosEnT H. MOHLENBROCK, JR. 


Continued studies of the flora of southern lllinois have re- 
sulted in the discovery of a number of species not previously re- 
ported from the state and of some species previously unknown 
in southern Illinois. 

PENSTEMON ARKANSANUS Pennell. The discovery of the 
Ozarkian species in an area where other members of this province 
have been found furthers one’s belief that southwestern Randolph 
County and northwestern Jackson County at one time harbored 
a considerable number of Ozark plants. Ranunculus harveyi, 
Talinum calycinum, Solidago buckleyi, and Pinus echinata are 
other southern species which occur in adjacent areas. In 
Missouri, Penstemon arkansanus is known from Madison, Ozark, 
Taney, Stone, Barry, Newton, and MeDonald Counties (Palmer 
and Steyermark, 1935). It also occurs in Arkansas and eastern 
Texas. The Illinois station which is on a rocky wooded hillside 
southwest of Ava in Jackson County (May 21, 1956; Mohlenbrock 
$808) extends the range of this species to the north and east. 
Penstemon arkansanus resembles P. pallidus in several respects, 
but the leaves of P. arkansanus are glabrous or nearly so and 
the flower parts are somewhat smaller. 

PorvaoNaTUM. In recent years, the literature concerning the 
occurrence of species of the genus Polygonatum in Illinois has 
become confusing. Jones in 1945 recognized P. biflorum (Walt.) 
Ell. and P. pubescens (Willd.) Pursh as occurring in Illinois. 
However, in 1950, Jones listed P. commutatum (Schult.) Dietr. 
and P. pubescens, placing the specimens previously included by 
him under P. biflorum into either P. commutatum or P. pubescens. 
This was on the basis that the true P. bzflorum of Elliott did not 
occur in Illinois. Fernald (1950) and Gleason (1952) substituted 
P. canaliculatum (Muhl.) Pursh for P. commutatum since Muhlen- 
berg's epithet was recorded before that of Schultes. However, 
in a fairly recent monograph on Polygonatum, Ownbey (1944) 
states that since not a single character in the description of 
Convallaria canaliculata Muhl. corresponds with the characters 
of the present Polygonatum commutatum, it is best to consider 
C. canaliculata as a synonym for P. biflorum. 


1958] Mohlenbrock, —Field and Herbarium Studies 203 


After collecting a specimen of what proved to be P. biflorum 
from Pope County, the author was led to study the problems 
concerning Polygonatum in Illinois. The following key serves 
to separate the three species that occur in Illinois. 


a. Leaves pilose on the nerves beneath..................... 1. P. pubescens. 

a. Leaves glabrous beneath... . 20.0.0... 000 eee b 
b. Leaves more or less clasping or sheathing at the base, the 
largest ones with over 100 nerves; perianth 17-20 mm. 

long, the lobes 5-7 mm. long.................... 2. P. commutatum. 
b. Leaves sessile at the base, the largest ones with less than 
100 nerves; perianth 10-17 mm. long, the lobes 3-4 mm. 

INE UU iN luo ea rer ther RB 3. P. biflorum. 


1. PoLYGONATUM PUBESCENS (Willd.) Pursh. In Illinois, this species 
grows in relatively moist, shaded woods in the northeast section of the 
state. It has been collected in Cook, DuPage, Kankakee, Lake, and 
Winnebago Counties, the collection from the latter county by Fell in 
1945 being the only one since 1914. 


2. PoLYGONATUM COMMUTATUM (Schult.) Dietr. This is the common 
species of Polygonatum, found throughout the state in a wide variety of 
habitats. 


3. PoLYGONATUM BIFLORUM (Walt.) Ell. Only three collections for this 
species have been seen from Illinois where it grows in upland woods. 
All the stations are in the extreme southern part of the state. SPECIMENS 
EXAMINED: Pope Co.: atop sandstone bluff, Belle Smythe Springs, April 
22, 1956, Mohlenbrock 8015; along Lusk Creek, May 21, 1952, Bailey & 
Swayne 2359. Hardin Co.: closed woods, Rock Creek valley, May 3, 1949, 
Bailey & Swayne 673. 


Tur NATIVE SPECIES OF Pyrus. E. J. Palmer in making his 
botanical reconaissance through southern Illinois in 1919 re- 
ported finding the narrow-leaved crab apple, Pyrus angustifolia, 
in Pope and Johnson Counties. These specimens, deposited in 
the Gray Herbarium and the herbarium of the Missouri Botan- 
ical Garden, have been considered merely as morphological varia- 
tions of Malus (Pyrus coronaria) by Jones (1945, 1950) and 
Jones, et al. (1955), and do not represent the species which Aiton 
described. In the fall of 1957, the author found specimens 
which match Aiton’s entity while collecting in a pin oak, 
overcup oak flat in Jackson County. Although Pyrus angusti- 
folia is predominantly a plant of the southeast, it does extend 
into Kentucky and southern Illinois. Pyrus lancifolia Rehder, 
a plant which somewhat resembles P. angustifolia but differs 


294 Rhodora [Vor. 60 


principally in leaf shape and flower and fruit size, is best treated 
as a variation of the more widespread P. coronaria L. Another 
species, P. ?oensis (Wood) Bailey, has characters which separate 
it more easily from the other taxa. 

The native members of the genus Pyrus, subgenus Malus 
in Illinois may be separated as follows: 


a. Calyx densely tomentose on the outside; leaves on vigorous 


shoots tomentose or villous at maturity. ...........0........ P. ioensis. 

a. Calyx glabrous or nearly so on the outside; leaves glabrous or 
villous along the veins beneath at maturity...............0.......... b 

b. Leaves on vigorous shoots more than half as broad as long, 
distinctly lobed...........iiiii sese eee 2. P. coronaria. 

b. Leaves on vigorous shoots less than half as broad as long, 
serrate or shallowly lobed... 20... 000000 00000 sss c 


c. Leaves of fertile branches acute or acuminate at the apex, 

more or less rounded at the base; pedicels 3.0-3.5 cm. long; 

petals 1.0-1.5 em. broad. .............. 2a. P. coronaria var. lancifolia. 
c. Leaves of fertile branches obtuse to subacute at the apex, 

cuneate at the base; pedicels 2.0-2.5 em. long; petals 0.5— 

1.0 em. broad... esses 3. P. angustifolia. 


1. Pyrus roENsIS (Wood) Bailey. The Iowa Crab Apple is common in 
and along the edges of oak-hickory woods throughout Illinois. It is a 
frequent intruder in the hill prairies along the bluffs of the Mississippl 
River. Specimens from Richland (Ridgway in 1928) and Lawrence 
(Eaton in 1901) Counties determined as Malus platycarpa var. hoopesii 
(Rehder) Rehder may represent hydrids between P. ioensis and some other 
species, but with our present knowledge, this entity is best treated as a 
morphological variant of P. ioensis. 


2. PYRUS coronaria L. The Wild Sweet Crab Apple is an occasional 
species of rather mesic woods in southern and eastern Illinois. Narrow- 
leaved forms may be segregated as: PYRUS CORONARIA L. var. LANCIFOLIA 
(Rehder) Fern. in Rhodora 49: 232. 1947. 

This variant is known through a few collections in southern Illinois now 
deposited in the Gray Herbarium from the counties of Jackson, Gallatin, 
and Pope. 


3. Pyrus ANGUSTIFOLIA Ait. This southern species has two known sta- 
tions in Illinois where it grows in bottomland woods. The leaves remain 
green far into November. Jackson Co.: in pin oak-overeup oak flats, 
eight miles southwest of Murphysboro, Mohlenbrock 8810. Hardin Co.: 
near Rosiclair, October 10, 1919, E. J. Palmer 17094. 


HELIANTHUS HIRSUTUS Raf. var. TRACHYPHYLLUS T. & G. 
This variety which differs from var. hirsutus in having internodes 
short-hispid and scabrous and leaves 2-3 em. broad can now 


Mohlenbrock,— Field and Herbarium Studies 295 


be added to the flora of Illinois on the basis of its collection in 
Jackson County: dry open woods, Lake Murphysboro Recreation 
Area, August 31, 1957, Mohlenbrock 8005. 

SMILAX GLAUCA Walt. var. LEUROPHYLLA Blake. "This smooth- 
leaved variety of lowland areas has been found for the first 
time in Illinois growing along the edge of LaRue Swamp in 
southwestern Illinois (Union County): edge of water, LaRue 
Swamp, three miles north of Wolf Lake, June 22, 1956, Mohlen- 
brock 7926. 

BoEHMERIA DRUMMONDIANA Wedd. This species of false 
nettle, often considered as a variety of B. cylindrica (L.) Sw., has 
been found in a boggy area in southern Illinois (1 mile north 
of Murphysboro, Mohlenbrock 756) where it is associated with 
Dryopteris thelypteris, Solidago patula, Eupatorium perfoliatum, 
and other moisture-loving plants. The narrower, harshly 
scabrous, short-petioled leaves distinguish this species from 
B. cylindrica. 

PrLEA IN ILLINOIS. In treating the species of Adicea Raf. 
( = Pilea) of northern and central United States, Lunell recognized 
five species, four of which he named for the first time in 1913. 
Of these, three species have now been found in Illinois. All 
Illinois specimens of P. opaca have been collected since the pub- 
lication of ‘‘Flora of Illinois" (Jones, 1950), and Fernald (1950) 
did not include this species in his manual. The following key 
separates these three species. 


a. Achenes green, averaging 1 mm. wide...................5. 1. P. pumila. 
a. Achenes black, averaging 1.5 mm. wide...............0. ee ee ee ee eeees b 
b. Plants leafy above the middle; seeds averaging 1.5 mm. 
| (0) 11 OER RA Den Mo UE LL ILI 2. P. fontana. 
b. Plants branching and leafy from near the base; seeds aver- 
aging.2.0mm-long...... x. ee 3. P. opaca. 


1. PrLEA PUMILA (L.) Gray. The common clearweed is abundant in moist 
situations throughout Illinois. Variation in the leaf margins occurs, 
leading to the naming of scarcely separable taxa such as var. deamii 
(Lunell) Fern. The light-colored fruit easily separates this species from 
the following two. 


2. PILEA FONTANA (Lunell) Rydb. This is the smallest species of Pilea. 
Lunell described it '4-8 em. altus", but most specimens observed fre- 
quently become 20 em. tall. Usually the teeth on the margin of the 
leaf are much lower than in either P. pumila or P. opaca. The two col- 


296 Rhodora [Vor. 60 


lections in the University of Illinois herbarium are the only ones known 
from Illinois. SPECIMENS EXAMINED: Grundy Co.: on rotting post in 
Illinois River, September 28, 1951, Ahles 4058. Vermilion Co.: on top 
of fill from a ditch in swamp, Muncie, October 2, 1950, Ahles 3395. 
3. Pinna opaca (Lunell) Rydb. This species also grows to a size larger 
than that ascribed to it in the original deseription. Lunell states that 
the species grows “10-30 em. altus". While most of our specimens fit 
this range, some attain a height of 60 em. The previously known records 
of P. opaca from Illinois are from Champaign, Cook, Iroquois, Piatt, 
and Vermilion Counties (collection data can be obtained by consulting 
Jones, et al., 1955). Now, Union County ean be added: low ground at 
base of limestone bluff, Pine Hills north of Wolf Lake, June 15, 1956, 
Mohlenbrock. 


LITHOSPERMUM CROCEUM Fern. This species is included by 
Jones, et al., (1955) under L. caroliniense (Walt. MacM. Re- 
gardless of which species one recognizes, the Jackson County 
collection is the first from southern Illinois. A summary of 
characters between L. croceum and L. caroliniense follows: 


L. croceum L. caroliniense 
Leaves over 30 on the stem Leaves 25 or less on the stem 
Leaves with papillose-based hairs Leaves without papillose-based hairs 
Braets closely overlapping Bracts often remote 
Calyx lobes keeled, papillose Calyx lobes flat, not papillose 
Veins of calyx anastomosing Veins of calyx free 


COLLECTION DATA: dry oak-hickory woods, Lake Murphysboro Recrea- 
tion Area, May 15, 1957, Mohlenbrock 8021. 


EUPHORBIA COMMUTATA Engelm. The wood spurge is a very 
attractive species in the early spring since the upper leaves and 
bracts are red while the lower leaves are green. In addition to 
its known range in seven of Illinois’ northern counties, it has 
been found now in Williamson County in the Devil's Kitchen 
area: mesie woods, nine miles southeast of Carbondale, March 
30, 1957, Mohlenbrock 8010. 

BROMUS BRIZAEFORMIS Fisch. & Mey. In searching through 
the herbarium at Southern Illinois University, a specimen of the 
rattlesnake chess collected in Washington County by French 
in 1872 was found, making the second known collection for this 
species in Illinois. It was collected previously from Richland 
County by Ridgway in 1902. 


1958] Bean et al.,— Report of Committee on Plant Distribution 297 


LITERATURE CITED 


FERNALD, M. L. 1950. Gray’s Manual of Botany. 

GLEASON, H. A. 1952. New Britton and Brown Illustrated Flora. 

JoNES, G. N. 1950. Flora of Illinois. 

————— ——, G. D. Futter, G. 8. WiwTERRINGER, H. E. AHLES, AND 
A. Frynn. 1955. Vascular Plants of Illinois. 

Ownsey, R. P. 1944. The Liliaceous Genus Polygonatum in North 
America. Ann. Missouri Bot. Gard. 31: 373-413. 

PALMER, E. J. AND J. A. STEYERMARK. 1935. An Annotated Catalogue 
of the Flowering Plants of Missouri. Ann. Missouri Bot. Gard. 22: 
375-758. 


TWELFTH REPORT OF THE COMMITTEE ON PLANT 
DISTRIBUTION 


The eleventh report included the Dicotyledoneae through 
Moraceae. The present report deals with the families from 
Cannabinaceae through Aizoaceae, taken in the order of the 
eighth edition of Gray’s Manual. Although the Polygonaceae 
were treated by Mr. Wm. P. Rich in 1902 (Rhodora IV-203), 
they are discussed again at this time. 

'The data for these reports have been compiled from the her- 
barium of the New England Botanieal Club and from the Gray 
Herbarium. 


PRELIMINARY Lists oF New ENGLAND PLANTS— XXXVII 


The sign + indicates that an herbarium specimen has been 
seen, the sign — that a reliable printed record has been found 
and the sign * is used for those plants which are not native in 
our flora. 


E A E "RA 
P Z > P [ae Q 
CANNABINACEAE 
*Cannabis sativa L. + + + + zi + 
*Humulus japonicus Sieb. & Zucc. + + + + + 
*Humulus Lupulus L. + + + + 3E 4 
URTICACEAE 
Boehmeria cylindrica (L.) Sw. -+ + ae B zu 4 
Boehmeria cylindrica var. Drummondiana Wedd. + + + = 
Laportea canadensis (L.) Wedd. + + + + E RD 
Parietaria floridana Nutt. + 
Parietaria pensylvanica Muhl. + + + + 
Pilea pumila (L.) Gray + + + I T + 


208 Rhodora 


[Vor. 60 


Urtica chamaedryoides Pursh 
*Urtica dioica L. 

Urtica gracilis Ait. 

Urtica procera Muhl. 

*Urtica urens L. 
Urtica viridis Rydb. 
SANTALACEAE 

Comandra Richardsiana Fern. 

Comandra umbellata (L.) Nutt. 

Geocaulon lividum (Richards.) Fern. 

LORANTHACEAE 

Arceuthobium pusillum Peck 

ARISTOLOCHIACEAE 
*Aristolochia Clematitis L. 
*Aristolochia durior Hill 
Aristolochia Serpentaria L. 
Asarum canadense L. 
Asarum canadense var. acuminatum Ashe 
Asarum canadense var. reflexum (Bickn.) Robins. 
POLYGONACEAE 
*Emex spinosa Campd. 
*Fagopyrum sagittatum Gilib. 
*Fagopyrum tataricum (L.) Gaertn. 
Oxyria digyna (L.) Hill 
Polygonella articulata (L.) Meisn. 
Polygonum achoreum Blake 
Polygonum allocarpum Blake 
*Polygonum amphibium L. f. terrestre (Leers) Fern. 
Polygonum amphibium var. stipulaceum 'Coleman) 
Fern. 

Polygonum amphibium var. stipulaceum f. fluitans 
(Eat.) Fern. 

Polygonum amphibium var. stipulaceum f. hirtuosum 
(Farw.) Fern. 

Polygonum amphibium var, stipulaceum f. simile Fern. 
Polygonum arifolium L. var pubescens (Keller) Fern. 
*Polygonum aviculare L. 

Polygonum aviculare var. littorale (Link) W. D. J. Koch 
*Polygonum aviculare var. vegetum Ledeb. 
*Polygonum Bistorta L. 

Polygonum Careyi Olney 
*Polygonum cespitosum Blume var.  longisetum 

(DeBruyn) Stewart 
Polygonum cilinode Michx. (including f. erectum (Peck) 
Fern.) 

Polygonum coccineum Muhl. 

Polygonum coccineum f. natans (Wieg.) Stanford 
*Polygonum Convolvulus L. 

*Polygonum Convolvulus var, subalatum Lej. & Court. 

Polygonum cristatum Engelm. & Gray 
*Polygonum cuspidatum Sieb. & Zucc. 

Polygonum Douglasii Greene 
*Polygonum dubium Stein 
Polygonum erectum L. 


++ 


++4+4+++ 


+ 
T 


+++ 44444 


++44 


+ 


+ +444 


+ ++ ++ ++ 


+ + +++ +++ 


++ 


++ ++ 


+ +1444 


m 


+ + +++ 


+ 


++4+4444 


++ +44+444+4+ 


++ 


+ +444 


++++ ++ 


— 


++ +++++ c +++++ 


++H+++++ 


+ 


1958] Bean et al.,— Report of Committee on Plant Distribution 299 


I 


Conn. 


+ | Me. 
N 
Vt 


Polygonum Fowleri Robins. 
Polygonum exsertum Small 4 4 -+ 
Polygonum glaucum Nutt. + + 
Polygonum Hydropiper L. (including var. projectum 
Stanford) + 
Polygonum hydropiperoides Michx. + 
Polygonum hydropiperoides f. strigosum (Small) 
Stanford 
Polygonum lapathifolium L. + 
*Polygonum lapathifolium var. prostratum Wimm. 
Polygonum lapathifolium var. salicifolium Sibth. + 
*Polygonum minus Huds. var. subcontinuum (Meisn.) 
Fern. 
*Polygonum nepalense Meisn. 
Polygonum opelousanum Riddell 
Polygonum opelousanum var. adenocalyx Stanford 
*Polygonum orientale L. 
Polygonum pensylvanicum L. 
Polygonum pensylvanicum var. laevigatum Fern. 
Polygonum pensylvanicum var. laevigatum f. albineum 
Farw. 
Polygonum pensylvanicum var. nesophilum Fern. 
*Polygonum Persicaria L. 
*Polygonum Persicaria var. ruderale (Salisb.) Meisn. 
*Polygonum polystachyum Wall. 
Polygonum prolificum (Small) Robins. 
Polygonum punctatum Ell. 
Polygonum punctatum var. leptostachyum (Meisn.) 
Small 
Polygonum punctatum var. parvum Vict. & Rousseau 
Polygonum puritanorum Fern. 
Polygonum ramosissimum Michx. 
Polygonum ramosissimum f. atlanticum Robins. 
Polygonum robustius (Small) Fern. 
*Polygonum sachalinense F. Schmidt 
Polygonum sagittatum L. 
Polygonum sagittatum f. chloranthum Fern. 
*Polygonum scabrum Moench 
Polygonum scandens L. 
Polygonum setaceum Baldw. var. interjectum Fern. 
Polygonum tenue Michx. 
Polygonum viviparum L. 
*Rheum Rhaponticum L. 
*Rumex Acetosa L. 
*Rumex Acetosella L. 
*Rumex alpinus L. 
Rumex altissimus Wood 
*Rumex crispus L. 
Rumex crispus X obtusifolius 
Rumex crispus X orbiculatus 
*Rumex domesticus Hartm. 
Rumex fenestratus Greene 
Rumex hastatulatus Baldw. 
*Rumex maritimus L. 


--- 


++ ++ 
++ 


+ 
— 


+ 
+ 
+ + +++ ++ + ++ ++ 


+++ +++++ 


++ +4 
+ ++ ++ 
l 

++ 


++i+++ + ++ 


++ + + 4+ 
+ + +4+4+14+4 


+ +4444 


++++ + + +++ 
++ ++ 


+ +++ + +4 


= 


++ +t++++++Ii+ ++t4t4+ +++++ 
+ 


+++ +++ +++ +H+++++ttt++t+ ++ttt++ +++++ + ++++ ++ 


300 Rhodora [Vor. 60 


Me. 
N. H. 
Vt. 
Mass. 


| 


+++ ++ ++ ++ +++ GRE 


Rumex maritimus var. fueginus (Phil.) Dusén + 
*Rumex mexicanus Meisn. (including R. triangulivalvis 

(Danser) Rech. f.) + 

*Rumex obtusifolius L. + 

Rumex orbiculatus Gray + 

T 

+ 


+++ + | Conn. 


Rumex pallidus Bigel. . 

Rumex persicarioides L. 

*Rumex Patientia L. 

*Rumex pulcher L. 

Rumex verticillatus L. 

Tovara virginiana (L.) Raf. 
CHENOPODIACEAE 

Atriplex arenaria Nutt. 

Atriplex glabriuscula Edmondston 
*Atriplex hortensis L. 

Atriplex patula L. 

Atriplex patula var. hastata (L.) Gray 

Atriplex patula var. littoralis (L.) Gray 
*Atriplex rosea L. 

*Axyris amaranthoides L. 

*Bassia hirsuta (L.) Aschers. 

*Beta vulgaris L. 

*Chenopodium album L. 

*Chenopodium ambrosioides L. 

*Chenopodium ambrosioides var. anthelminticum (L.) 
Gray 

*Chenopodium Bonus-Henricus L. 

Chenopodium Boscianum Moq. 
*Chenopodium Botrys L. 

Chenopodium capitatum (L.) Aschers. 
*Chenopodium carinatum R. Br. 
*Chenopodium foliosum (Moench) Aschers. 
*Chenopodium glaucum L. 

*Chenopodium graveolens Lag. & Rodr. 

Chenopodium humile Hook. 

Chenopodium hybridum L. var. gigantospermum 

(Aellen) Rouleau 
*Chenopodium incanum (S. Wats.) Heller 
*Chenopodium lanceolatum Muhl. 

Chenopodium leptophyllum Nutt. 
*Chenopodium murale L. 

*Chenopodium paganum Reichenb. 
*Chenopodium polyspermum L. 

Chenopodium rubrum L. 
*Chenopodium urbicum L. 
*Corispermum hyssopifolium L. 

*Cycloloma atriplicifolium (Spreng.) Coult. 
*Kochia Scoparia (L.) Roth 
*Kochia Sieversiana (Pall.) C. A. Mey. 
*Monolepis Nuttalliana (R. & S.) Greene 
*Roubieva multifida (L.) Moq. 

Salicornia Bigelovii Torr. 

Salicornia europaea L. 

Salicornia europaea var. prostrata (Pall.) Fern. 


++ + 4444+ 
+++44+44+4+ 4+ 


++4+4+4+ ++ + 


++ + +++ 4 
+++ + +++ ++ 4 
— 


++++ 
++++ 
+++ 
+++ ++ 
+++ ++) 


I +t+4+t+4+4¢4+5+ +41 
+ ++ + ++ + + 
| +) + + + + 
+H++++++4+4+ +++ +  Ft+Htt+tt+e F444 44444441 
++++ + 
++ ++ I++ + 


++ 
++ 
++ 


+++ + 


1958] Bean et al.,—Report of Committee on Plant Distribution 301 


| 


i+ +++4++++ +++] Me 


Salicornia europaea var. simplex (Pursh) Fern. 
Salicornia virginica L. 
Salsola Kali L. 
Salsola Kali var. caroliniana (Walt.) Nutt. 
*Salsola Kali var. tenuifolia Tausch 
*Spinacea glabra Mill. 
*Spinacea oleracea L. 
Suaeda americana (Pers.) Fern. 
Suaeda linearis (Ell.) Moq. 
Suaeda maritima (L.) Dumort. 
Suaeda Richii Fern. 
AMARANTHACEAE 
*Acnida altissima Riddell 
*Acnida altissima var. prostrata (Uline & Bray) Fern. 
*Acnida altissima var. subnuda (S. Wats.) Fern. 
Acnida cannabina L. 
*Acnida tamariscina (Nutt.) Wood 
Amaranthus albus L. 
*Amaranthus ascendens Loisel. 
*Amaranthus caudatus L. 
*Amaranthus cruentus L. 
*Amaranthus deflexus L. 
*Amaranthus graecizans L. + + + 
*Amaranthus hybridus L. (including f. hypochondriacus 
(L.) Robins.) + ae ES 
*Amaranthus lividus L. 
*Amaranthus Palmeri S. Wats. 
*Amaranthus Powellii S. Wats. — — 
Amaranthus pumilus Raf. 
*Amaranthus retroflexus L. + + + 
*Amaranthus spinosus L. — 
*Amaranthus viridis L. 
*Celosia argentea L. var. cristata (L.) Ktze. 
*Froelichia arizonica Thornber 
NYCTAGINACEAE 
*Mirabilis hirsuta (Pursh) MacM. + 
*Mirabilis Jalapa L. 
*Mirabilis linearis (Pursh) Heimerl 
*Mirabilis nyctaginea (Michx.) MacM. 
PHYTOLACCACEAE 
Phytolacca americana L. 
AIZOACEAE 
*Mollugo verticillata L. E 
*Tetragonia expansa Murr. 


++++ | Conn. 


++ + + +44 va 
PEI 


++ + +t++++|RL 


+++ 
+ 
ar a ete 
+ +++++ ++++ | Mass, 
+ ++ı+ ++ 


++ + 
++ + + +1 


+ + +++ F444 44444444 
+ 
+ 


++ + ++++ 


++ + + 


With the exception of the introduced species the geographical 
areas are the same as in previous reports. The groups repre- 
sented are in marked contrast to those treated in the eleventh 
report. While the species in the latter were with one exception 
woody, in the present one there are no trees or shrubs. In the 
last report there were no strictly maritime species, while here 


302 Rhodora [Vor. 60 


there is a large representation of such plants. Another notable 
difference is that nearly fifty per cent of the forms here treated 
are not native to New England, but are introduced, naturalized 
or adventive. Several subgroups have consequently been 
adopted in this category. When only one, or at most five or 
six specimens representing a given form have been found, it has 
seemed best to designate such a plant as “local” even though in 
the area covered by Gray’s Manual it may be distinctly northern 
or southern and in some regions common. In New England 
there are not enough stations to show any definite distributional 
patterns in such cases. 


I. GENERALLY DISTRIBUTED.—Urtica procera, Polygonum amphibium 
rar. stipulaceum f. fluitans, P. coccineum, P. coccineum f. natans, P. 
Hydropiper (including var. projectum), P. sagittatum, Rumex orbiculatus. 

Ia. GENERAL, EXCEPT MAINE COAST EAST OF KENNEBEC RIVER.—Pilea 
pumila, Polygonum pensylvanicum var. laevigatum. 

Ib. GENERAL, EXCEPT THE COAST OF WASHINGTON COUNTY, MAINE.— 
Polygonum lapathifolium, P. punctatum (including var. leptostachyum), 
Amaranthus albus. 

Ic. GENERAL, EXCEPT CAPE COD.—Polygonum cilinode (including var. 
erectum). 

Id. GENERAL, EXCEPT CAPE COD AND MAINE COAST EAST OF KENNEBEC 
RIVER.—Urtica gracilis. While Urtica gracilis seems to belong to this 
group, it does not occur on the coast north of Massachusetts and is very 
rare in New Hampshire. 

Ie. GENERAL, EXCEPT CAPE COD AND WASHINGTON COUNTY, MAINE— 
Laportea canadensis, Polygonum amphibium var. stipulaceum, Chenopodium 
hybridum var. gigantospermum. 

IIA. NORTHERN—NUMEROUS STATIONS SOUTH OF 43°.—Arceuthobium 
pusillum, Polygonum lapathifolium var. salicifolium, Chenopodium capita- 
tum. 

Polygonum lapathifolium var. salicifolium occurs infrequently and 
there are no stations on the coast north of Massachusetts. Chenopodium 
capitatum is absent from the coast and infrequent elsewhere. 

IIB. NORTHERN—NOT OR NOT MUCH SOUTH OF 43°.— Urtica viridis, 
Geocaulon lividum, Polygonum Douglasii, Rumex fenestratus. 

Urtica viridis is known only from the Maine coast east of the Kennebee 
River. Polygonum Douglasii is somewhat local and the stations for this 
species lie between 43? and 45° except on the coast. Rumex fenestratus 
is found in Washington County only. 

III. ALPINE.—Oxzyria digyna is found on the White Mountains only. 
Polygonum viviparum occurs also on Mt. Mansfield and Mt. Katahdin. 

IV. CAPE COD BUT NOT NORTHERN MAINE—GENERAL IN MAINE SOUTH 
or 45°.—Boehmeria cylindrica, Comandra umbellata, Polygonum arifolium 
var. pubescens, P. Careyi, P. hydropiperoides, Chenopodium leptophyllum. 


1958] Bean et al.,— Report of Committee on Plant Distribution 303 


Boehmeria cylindrica does not occur east of the Penobscot River. 
Polygonum arifolium var. pubescens is not found either in northern New 
Hampshire or northern Vermont. Chenopodium leptophyllum is infre- 
quent and there are no stations for it in Vermont. It occurs in New 
Hampshire only in the Androscoggin Valley. 

V. NEITHER CAPE COD NOR NORTHERN MAINE, NOT IN WASHINGTON 
county.—Polygonum erectum, Chenopodium Boscianum. 

While Chenopodium Boscianum does not occur on Cape Cod, it is present 
on Nantucket and is not found north of southwestern Maine (S. Berwick). 

VIA. CHIEFLY THE THREE SOUTHERN STATES, BOTH CAPE COD AND 
WESTERN MASSACHUSETTS.—Polygonnm cristatum, P. tenue, Phytolacca 
americana. 

VIB. CHIEFLY THE THREE SOUTHERN STATES, CAPE COD BUT NOT WEST- 
ERN MASSACHUSETTS.—Polygonella articulata, Polygonum scandens. 

Polygonella articulata seems to avoid the rich soils of western New 
England. It is found along the Maine coast to the Kennebec River. It 
is frequent along railroads and in sandy soil. Polygonum scandens also 
avoids the rich soils of western Vermont and Massachusetts. 

VIC. CHIEFLY THE THREE SOUTHERN STATES, NEITHER CAPE COD NOR 
WESTERN MASSACHUSETTS.—Polygonum robustius. 

VID. SOUTHWESTERN NEW ENGLAND ONLY.—Aristolochia Serpentaria. 

VII. COASTAL PLAIN.—Boehmeria cylindrica var. Drummondiana, Poly- 
gonum opelousanum, P. opelousanum var. adenocalyx, P. pensylvanicum, 
P. pensylvanicum var. nesophilum, P. puritanorum, P. setaceum var. 
interjectum, Rumex verticillatus. 

While Boehmeria cylindrica var. Drummondiana seems to fit this cata- 
gory it has two stations far outside the coastal plain, one at Beverly, 
Massachusetts, the other at Dorset, Vermont. Polygonum puritanorum 
has one isolated station at Waterford, Maine. 

VIII. CALCICOLOUS—CHIEFLY WEST OF THE CONNECTICUT RIVER IN 
SOUTH; IN EAST MOSTLY NORTH OF 45°.— Parietaria pensylvanica, Comandra 
Richardsiana, Asarum canadense, A. canadense var. acuminatum, A. 
canadense var. reflexum, Polygonum achoreum, Tovara virginiana. 

Parietaria pensylvanica occurs in western New England, on the Cran- 
berry Islands, Maine and locally at three seemingly indigenous stations 
near Boston. Comandra Richardsiana is restricted to the northern Lake 
Champlain region. Asarum canadense also occurs on soils overlying basic 
rocks in eastern Massachusetts. Asarum canadense var. reflexum is 
restricted to southwestern Connecticut, while Asarum canadense var. 
acuminatum occurs west of the Connecticut River only. Polygonum 
achoreum is found at but four stations: Castle Hill and Caribou in Aroos- 
took County, Maine, and Grand Isle and South Burlington in Vermont. 
Tovara virginiana is frequent in western New England and also occurs in 
seemingly indigenous locations near Boston and at two stations in Rhode 
Island. 

IXA. MARITIME HALOPHYTES—IN VICINITY OF COAST, NO INLAND STA- 
TIONS.—This category is further divided into three subgroups— 


304 Rhodora [Vor. 60 


a. GENERAL.—Polygonum prolificum, P. ramosissimum var. atlanticum, 
Atriplex glabriuscula, A. patula var. littoralis, Salicornia europaea, Suaeda 
linearis, S. maritima. 

b. NORTHERN.— The southern limit is given for each species: Polygonum 
allocarpum extends to Cape Elizabeth, Maine, Rumex pallidus to Massa- 
chusetts Bay with one station at Nantucket, R. persicarioides from Cape 
Ann to Block Island only. Chenopodium rubrum occurs chiefly in New 
Hampshire and Massachusetts with one station each in Maine and Con- 
necticut. Salicornia europaea var. prostrata extends to Penobscot Bay, 
Suaeda americana to Cape Ann and S. Richii to Massachusetts Bay. 

c. SOUTHERN.—The northern limit of each species is stated: Polygonum 
ersertum extends to S. Berwick, Maine, P. glaucum to Massachusetts 
Bay, Atriplex arenaria to Hampton, New Hampshire, Salicornia Bigelovii 
to Cape Elizabeth, Maine, S. europaea var. simplex to Hampton, New 
Hampshire, S. virginica to Wells, Maine and Acnida cannabina to Casco 
Bay. 

IXB. MARITIME (NOT NECESSARILY HALOPHYTIC) WITH INLAND STA- 
TIONS.—Polygonum aviculare var. littorale, P. ramosissimum, Rumex 
maritimus var. fueginus, Atriplex patula, A. patula var. hastata, Salsola 
Kali, S. Kali var. caroliniana. 

Salsola Kali var. caroliniana is absent from Maine, New Hampshire 
and Vermont. 

X. ESTUARINE.—Polygonum punctatum var. parvum, P. sagittatum var. 
chloranthum. 

XI. INTRODUCED SPECIES—GENERAL.—Humulus Lupulus, Fagopyrum 
sagittatum, Polygonum aviculare, P. aviculare var. vegetum, P. Convolvulus, 
P. Convolvulus var. subalatum, P. Persicaria, Rumex Acetosella, R. crispus, 
R. domesticus, R. mexicanus (including R. triangulivalvis), R. obtusifolius, 
Chenopodium album, C. lanceolatum, Salsola Kali var. tenuifolia, Amaran- 
thus retroflexus, Mollugo verticillata. 

Polygonum Convolvulus is absent from northern Maine. Neither 
Chenopodium lanceolatum nor Salsola Kali var. tenuifolia occurs north of 
45°. 

XIa. INTRODUCED SPECIES—NEITHER CAPE COD NOR NORTHERN MAINE.— 
These species occur very sparingly in the three northern states as shown 
by the following table: 


Number of stations 


Me. N.H. Vt 
Cannabis sativa 3 2 5 
Humulus japonicus 3 0 1 
Urtica dioica 3 1 1 
Urtica urens 1 1 1 
Polygonum orientale 2 2 1 
Rumex Acetosa 5 0 5 
Rumex altissimus 2 1 1 
Chenopodium Botrys 3 4 10 
Chenopodium urbicum 1 2 0 
Acnida altissima 1 4 


1958] Bean et al.,—Report of Committee on Flant Distribution 305 


XIb. INTRODUCED SPECIES WITH NORTHERN TENDENCIES.—Fagopyrum 
tataricum, Polygonum scabrum, Rumex alpinus. 

Rumex alpinus has been recorded at only two stations: Berwick in 
southern Maine and Wilmington in southern Vermont. 

XIc. INTRODUCED SPECIES WITH SOUTHERN TENDENCIES—CHIEFLY 
SOUTH oF 43°.—Polygonum cuspidatum, Chenopodium ambrosioides, C. 
ambrosioides var. anthelminticum, C. carinatum, C. graveolens, C. murale, 
Kochia Scoparia, Amaranthus caudatus, A. cruentus, A. graecizans, A. 
hybridus (including f. hypochondriacus), A. lividus, A. Palmeri, A. Powellii, 
A. spinosus, Celosia argentea var. cristata, Mirabilis hirsuta, M . nyctaginea. 

In 1902, no stations for Polygonum cuspidatum were known in either 
Vermont or Rhode Island. Chenopodium carinatum is confined to outer 
Cape Cod except for three inland stations on ballast or wool waste. 

XId. INTRODUCED SPECIES—SPORADIC.—Polygonum amphibium f. ter- 
restre, P. Persicaria var. ruderale, P. sachalinense, Rumex Patientia, Bassia 
hirsuta, Chenopodium glaucum, C. paganum. 

Bassia hirsuta is chiefly maritime with one station at Westford, Massa- 
chusetts. Chenopodium paganum is absent from western Massachusetts 
and Connecticut. 

XIe. INTRODUCED SPECIES—LOCAL.—Aristolochia Clematitis, A. durior, 
Emex spinosa, Polygonum Bistorta, P. cespitosum var. longisetum, P. 
dubium, P. lapathifolium var. prostratum, P. minus var. subcontinuum, P. 
nepalense, P. polystachyum, Rheum Rhaponticum, Rumex maritimus, R. 
pulcher, Atriplex hortensis, A. rosea, Axyris amaranthoides, Beta vulgaris, 
Chenopodium Bonus-Henricus, C. foliosum, C. incanum, C. polyspermum, 
Corispermum hyssopifolium, Cycloloma atriplicifolium, Kochia Sieversiana, 
Monolepis Nuttalliana, Roubieva multifida, Spinacea glabra, S. oleracea, 
Acnida altissima var. subnuda, A. tamariscina, Amaranthus ascendens, A. 
caudatus, A. deflexus, A. viridis, Froelichia arizonica, Mirabilis Jalapa, 
M. linearis, Tetragonia expansa. 

XII. MISCELLANEOUS.—Parietaria floridana, Urtica | chamaedryoides, 
Polygonum Fowleri, Rumex hastatulatus, Chenopodium humile. 

Parietaria floridana is represented in New England by a single specimen 
from Pawtuckaway Mt., Rockingham County, N. H. Urtica cha- 
maedryoides is reported in the eighth edition of Gray's Manual to be ad- 
ventive north to Massachusetts, but there is no specimen from New 
England in the two herbaria consulted. Polygonum Fowleri is repre- 
sented by a single specimen in the New England Club herbarium from 
Isle au Haut. Other collections originally identified as P. Fowleri are 
now segregated as P. allocarpum. Rumex hastatulus was credited by 
Mr. Wm. P. Rich in Rhodora IV-205 to a single station at Salisbury 
Beach, Massachusetts. This specimen is now in the Gray Herbarium. 
During the fifty-six years only one other specimen has been collected: 
at Melrose, Massachusetts. Chenopodium humile is known from one 
station only in New England at Brunswick, Maine.—R. C. Bran, A. F. 
Hirn AND R. J. EATON. 


306 Rhodora [Vor. 60 


NorE ON THE DISTINCTION BETWEEN THE BROAD- AND 
NARROW-LEAVED ANTENNARIAS OF MriNNESOTA.— The broad- 
leaved and narrow-leaved Antennarias of the Gray's Manual 
range in the United States are customarily separated in keys on 
the basis of leaf size and nervation, with the size-ranges over- 


- 


lapping and the nerves varying in number from 3 to 7 in the 
broad-leaved plants to 1 or 3 in the narrow-leaved ones. Usually 
the nerves are said to be “prominent” in the former types and, 
when 3, “obscure” in the latter. Possibly this separation serves 
well enough for plants over most of the range, but for Minnesota 
plants it is unsatisfactory. Most of our narrow-leaved plants 
have 3 nerves, and these are often prominent. Close observation 
shows differences that make the separation more certain, par- 
ticularly for the beginning student. The following dichotomy 
distinguishes reasonably well between our broad-leaved plants 
(A. fallax, A. munda, and A. plantaginifolia of the 8th edition of 
Gray's Manual) and our narrow-leaved ones (4. aprica, A. 
campestris, A. canadensis, A. microphylla, A. neglecta, A. neo- 
dioica, and A. petaloidea of the 8th edition): 


Largest basal leaves 12-50 mm wide, 30-100 mm long, with 3-7 
lengthwise nerves at base of blade that are prominent beneath, 
the two (main) laterals nearest the midrib each located nearly 
always from 14 to 15 of the way from the midrib to the margin 
where farthest from the margin, usually extending past the 
broadest point of the blade and converging toward and nearly 
reaching the leaf tip, additional nerves if any branching mostly 
from the outer sides of the two laterals mentioned; smaller 
leaves and those not fully expanded often with lateral nerves 
as described below; largest blades tending to be widest be- 
tween 14 and 24 of the way to the tip; leaves of the flowering 
stem acute to bristle-tipped, without scarious appendages 

broad-leaved plants. 

Largest basal leaves 3-21 mm wide, 5-55 mm long, with 1 or 3 
prominent to obscure lengthwise nerves at base of blade, very 
rarely with a faint additional nerve next one or both margins, 
when 3- or more-nerved the two main laterals each located 
from nearly 24 to 34 of the way from the midrib to the margin 
in the broader part of the blade, usually extending at most 
shortly beyond the broadest point, rarely converging toward 
and nearly reaching the leaf tip, additional nerves if any 
branching mostly from the midrib; largest blades tending to be 
widest between 25 and 45 of the way to the tip; leaves of the 
flowering stem as above or the upper third of them each with a 
thin flattened sometimes inrolled scarious appendage 1.5-2.5 
mm. long by 0.4-0.9 mm wide. ............00 00000 ee narrow-leaved plants. 


—.THomas MORLEY, UNIVERSITY OF MINNESOTA. 


1958] Hermann,—Euphrasia micrantha 307 


EUPHRASIA MICRANTHA New To NomrH America.—Mount 
Greylock, a spur of the Taconic Range in Berkshire County, 
northwestern Massachusetts and the highest point (3491 ft. 
elevation) in the state, has already received considerable notice 
in the pages of this journal because of its botanical attractions. 
These have been revealed chiefly through the explorations of 
Dr. A. LeRoy Andrews, beginning with an account of the 
orchids of the region in 1900 and including a series of papers on 
the bryophytes extending up to 1957. Particularly noteworthy 
in the flora of the mountain is the subalpine element which, as 
might be expected, is markedly similar to that of the Green 
Mountains of Vermont. 

Early in the evening of August 26, 1957 the writer, returning 
from the Adirondacks, made a hurried drive to the summit of 
Greylock via the excellent road leading from North Adams, on 
the chance of getting in some botanizing before dusk set in. 
The most striking plant on the exposed rocky summit at this 
date proved to be a Euphrasia (F. J. Hermann 14075, specimens 
in Gray Herbarium, U. S. National Herbarium, N. Y. Botanical 
Garden Herbarium and Cambridge University Herbarium, Eng- 
land) occurring in extensive beds, its violet corollas vividly 
coloring the sparsely grassy stretches surrounding the war me- 
morial beacon which has become a popular tourist attraction. 

Later attempts to identify the plant in current floras were un- 
successful. None of the ten species included in Gray's Manual 
would accommodate its combination of blunt bract-teeth and 
almost solid violet corolla, nor could it be matched with any of 
the American collections in the U. 8. National Herbarium. A 
specimen sent to Kew was kindly referred by Dr. G. Taylor to 
Mr. P. F. Yeo, at Cambridge University Botanic Garden, who 
with Mr. P. D. Sell is at present engaged in a revision of the 
genus in North America. Mr. Yeo has referred the collection to 
the European Euphrasia micrantha Reichb., and very kindly 
sent European material of the species for comparison. As Mr. 
Yeo pointed out the plants from Mt. Greylock are not typical 
of the prevalent form of the species in Europe, differing chiefly 
in having very short floral internodes. In this respect it is very 
similar, however, to specimens collected by Mr. Yeo from cliff- 
top heaths in Brittany, so that it seems not unlikely that the 
shortening of the nodes is an ecological response. 


308 Rhodora [ Vou. 60 


The range of Euphrasia micrantha, which occurs throughout 
the British Isles according to Clapham, Tutin and Warburg in 
their Flora of the British Isles (1952), is given by H. W. Pugsley, 
in his Revision of the British Euphrasias (Journ. Linn. Soc. Bot. 
48:488. 1930), as “northwest Europe, including Scandinavia, 
Germany, Bohemia, and a large part of France”. The American 
species most nearly related to it is E. disjuncta Fern. & Wieg., 
of Labrador, Newfoundland and northern Maine (western plants 
formerly referred to E. disjuncta are really E. subarctica Raup, 
according to Hultén), which is also referred by Pugsley (Journ. 
Bot. 74:287. 1936) to the Series Latifoliae of the Subsect. Ciliatae, 
Sect. Semicalcaratae. Although characterized by blunt bract- 
teeth in common with E. micrantha, it is very unlike that plant 
in general aspect, the spikes being widely interrupted (lower 
bracts 2-5 cm. apart), and is readily distinguished from the 
Kuropean species by its larger, much broader and pubescent 
leaves and bracts. 

Although the occurrence of this European species in well- 
established colonies on a New England mountain top was un- 
expected, it does not seem improbable that it was introduced 
there in view of the fact that the number of visitors at the locality 
must run into many thousands each year, and that not a few 
other exotic plants have turned up in much less frequented 
areas.—F. J. HERMANN, CROPS RESEARCH DIVISION, A.R.S., U.S. 
DEPARTMENT OF AGRICULTURE, BELTSVILLE, MARYLAND. 


Volume 60, no. 718, including pages 265-288, was published 11 November, 
1958. 


QW R 


EFERENCE LIBRARY 


de 
R qq? 


Dodora 


JOURNAL OF THE 
NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
CARROLL EMORY WOOD, JR | 

IVAN MACKENZIE LAMB 


Vol. 60 December, 1958 No. 720 
CONTENTS: 
The Hybrid Ragweed, Ambrosia artemisiifolia X trifida. W. H. 
Wagner UTS 0 A a a aa eo cete aN eE S 309 
Some Interesting Records from Eastern Massachusetts. Richard 
PE ATN Sea cl T E IE eda sae 5 Ne IA 316 
Cabomba caroliniana Grows in Worcester County, Massachusetts. 
won N. Gales.. ce ae ee aie e a 321 
Further New Records of Myrtaceae from Jamaica. George R. 
Ptootor OVI c ve 6 5 cr ORDINI a ss re ee bas 323 
Typification of the Genus Forestiera (Oleaceae). Kenneth A. 
Wilson oe ie ERR es bee cc eee 327 
A New Rhododendron Station in Maine. Radcliffe Pike and 
AR. Hagan 1: e a cee a aa Ses SR MIS 328 
The Type of Vaucheria compacta var. koksoakensis, John L. 
Bim and Robert: T: W3tce EL Tie ciet ee 329 
iu. ii Dem CE ME PIT SEA LLENAS ES 330 
Index to volime:.00 .— 7... E CE er err cs re 331 


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Vol. 60 December, 1958 No. 720 


THE HYBRID RAGWEED, AMBROSIA ARTEMISIIFOLIA 
x TRIFIDA 


W. H. WAGNER, Jn.! 


One of the more striking of all weed hybrids is certainly the 
cross between the “common ragweed” and the “giant ragweed” — 
Ambrosia artemisiifolia X trifida. The parents differ strongly 
in habit, stature, leaf-form, and fruits, producing in their cross a 
distinctive plant. During recent studies of the parental species 
I have encountered occasional naturally occurring examples of 
this rare cross, and I wish to report here some observations not 
previously recorded. 

The hybrid between our two most pernicious of hayfever weeds 
was first discovered and deseribed by Wylie in 1915. Subsequent 
authors for the most part seem not to have noticed this early 
paper. A single plant was found during the summer of 1914 on a 
roadside about 2 miles north of Iowa City, Iowa. It produced 
flowers in abundance but no seeds were formed, although Wylie 
observed the plant closely until autumn. It is significant that 
none of the later writers has reported seeds (or fruits) either. 
The leaves of the new hybrid were described as ‘“‘distinctly 
different from either of the supposed parents, though inter- 
mediate between them in a general way,” and one of the leaves 
was figured (op. cit., pl. 21) together with comparable leaves of 
the parents. 

Since Wylie’s original description, various observations on 
Ambrosia artemisiifolia X trifida, both from North America 
and Europe, have been published. However, in his revision of 


1 Publication No. 10 on Atmospheric Pollution by Aeroallergens under research grant No. 
E-1379(C) from the National Institute of Allergy and Infectious Diseases, Public Health 
Service. 


310 Rhodora [ Von. 60 


Ambrosia in North American Flora Rydberg in 1922 did not men- 
tion Wylie's plant, including only another hybrid ragweed, A. 
bidentata X trifida. K. L. Jones (1943) did cite Wylie's original 
report and also recorded another example of the wild hybrid 
which was found in 1936 along the banks of the Huron River 
near Ann Arbor, Michigan. Jones, in genetic experiments, also 
created the hybrid under artificial conditions by growing the all- 
pistillate form of A. artemisiifolia together with pollinating 
plants of A. trifida. Of the seven hybrid plants that he secured, 
two inherited the all-pistillate condition of the artemisiifolia 
female parent, four were intergrades, and one was normally 
monoecious. In cytological studies of the cross, he determined 
that the 12 chromosomes in the haploid set of A. trifida are 
recognizably larger than the 18 from A. artemisiifolia. Chromo- 
some pairing was found to be irregular and there were numerous 
univalents. Such pairing as occurred involved almost entirely 
chromosomes of similar size and therefore were assumed to have 
been contributed by the same parent. Stebbins (1945) called 
attention to this as a striking example of pairing between chromo- 
somes contributed by the same parent in species hybrids involv- 
ing polyploidy. 

Rouleau in 1944 redescribed Ambrosia artemisiifolia X trifida 
on the basis of a collection made in Canada in 1935, and gave it 
the binomial “X Ambrosia Helenae," in honor of Hélène Boulé, 
for whom Ile Sainte-Héléne, where the hybrid was discovered 
near Montreal, was also named. In 1939, Ballais found the 
eross at Cauderan (Giroude), France, where the parent species 
have been introduced (Lawalrée, 1947). In 1950 the inter- 
mediate was taken at Urbana, Champaign Co., Illinois, by 
Marilyn L. Briggs (Jones and Fuller, 1955). 

Although we now have records in our notes only from Iowa, 
Illinois, Oklahoma, Michigan, Quebec, and France, Ambrosia 
artemisiifolia X trifida will probably be detected from time to 
time wherever the two parents grow together in ruderal habitats. 
In Michigan, as mentioned previously (Wagner & Beals, 1958), 
this ragweed cross is extremely rare. Such few specimens as 
we have found during 1956, 1957, and 1958 have been in much 
disturbed habitats, along the sides of newly built roads and on 
construction sites in and around Ann Arbor. 


1958] Wagner, —Hybrid Ragweed 311 


Despite its somewhat unique appearance the hybrid may be 
passed unnoticed, or may even be confused with other species. 
There are probably unrecognized specimens in various herbaria. 
To illustrate, in the herbarium of Tulane University I found in 
1958 an example of this ragweed cross from Pawnee, Oklahoma 
(August, 1933, C. F. Coffman) that was identified as Ambrosia 
psilostachya, the "perennial" or “western ragweed.” Another 
specimen reported for the first time here was found in 1957 in 
the much-used herbarium of the University of Michigan Bio- 
logical Station. The latter (East Cheboygan, Michigan, July 
30, 1933, J. H. Ehlers 5346) bore the identification A. artemisti- 
folia, probably because on casual inspection it appeared like a 
very coarse individual of the common ragweed. 

Part of the difficulty of recognizing A. artemisiifolia X trifida 
surely results from the rather extraordinary variation in the suc- 
cessive leaves produced from spring through fall. The earlier 
leaves of the season are roughly like over-sized leaves of A. 
artemisiifolia, but the later leaves formed on the main and lateral 
axes are not comparable directly to those of either parent, having 
rather large, but narrowly three-lobed or simple blades. By late 
summer and fall, all the spring leaves have normally fallen or 
completely dried up so that only the more simple leaf types of the 
distal parts of the plant remain. At this stage the plant looks 
something like a narrow-leaved form of A. trifida. 

To attempt to show all the normal foliar variations of this 
hybrid I decided to make observations of a single, as nearly 
average as possible, plant—from cotyledons to highest braet. 
To accomplish this it was necessary to find a natural hybrid 
very early in its growth, so that all the first leaves would be 
present and intact. On May 18, 1958, a specimen was found 
growing with the parents on a construction site at the University 
Hospital, Ann Arbor. With the assistance of Dr. John M. 
Sheldon this plant was extracted from a difficult position in a 
crack along the sidewalk with as little injury as possible, and it 
was turned over for cultivation to Mr. Walter F. Kleinschmidt, 
Superintendent of the U. M. Botanical Gardens, who kept it 
under approximately normal conditions. A sample leaf was re- 
moved from each node just before it dried and fell off naturally. 
The plant grew to 50 inches in height and seemed to be of fairly 


312 Rhodora | VoL. 60 


average growth for this cross in Michigan, neither as small as 
Ehlers’ specimen from Cheboygan, nor as large as the giant speci- 
men grown under more luxuriant artificial conditions the previous 
year. "The results from this “pruning” of the old leaves of suc- 
cessive nodes are shown in Figure 1. 

The seedlings of the parent species have been described by 
Rowlee (1893) and Kummer (1951). In over-all stature the 
hybrid seedling is closer to A. trifida than A. artemisiifolia. 
The cotyledons and the first foliage leaves of the cross are readily 
distinguishable from the corresponding leaves of A. trzfida. On 
the fourth to eighth nodes the leaves look somewhat like ex- 
tremely gross leaves of A. artemisiifolia and their margins are 
lobed and toothed (fig. 1, especially the second row). This is 
the leaf type depicted by Wylie (op. cit., pl. 21). By the time the 
sixth pair of foliage leaves has appeared, there are already lateral 
branches growing from all except the lowermost nodes, a feature 
especially characteristic of A. artemisiifolia (Rowlee, op. cit.). 
At approximately the middle level of the plant, the opposite leaf 
arrangement gives way to alternate, in this respect also as in 
A. artemisiifolia rather than A. trifida which has opposite branch- 
ing throughout. The leaves in the upper half of the plant tend 
to be progressively simpler in structure and finally, in the top 
five nodes, to become narrowly three-lobed to nearly simple, the 
margins now practically entire. The leaves illustrated by K. L. 
Jones (op. cit., figs. 2, 4, 5) are like the intermediate types found 
in the middle of the plant (fig. 1, third row). 

The leaves produced on the lateral branches of this “average” 
hybrid are considerably smaller than those along the main axis. 
The three leaves (fig. 1, inset) from a branch at the sixth node 
(shown by the arrow) illustrate the most common types of leaves 
on the lateral branches. In extremely vigorous, large and much- 
branched specimens of A. artemisiifolia X trifida, however, the 
lateral branches become more like the top half of the main axis 
of our average plant. This unusually large form is shown by the 
specimen of Rouleau (op. cit., fig. 3), and is represented in our 
collections by Wagner 8461 (Micu) from near Ann Arbor Airport 
which was grown to exceptionally large size in the U. M. Botani- 
cal Gardens in 1957. 

Fruits have apparently not previously been found in Ambrosia 


1958] Wagner, —Hybrid Ragweed 313 


FiG. 1. Leaf variations of Ambrosia artemisiifolia X trifida. | Lower left: Major branches of 


"average" plant 50 inches tall, showing successive nodes where leaves were removed. Sil- 
houettes (to scale) of successive leaves arranged left to right—Bottom row, nodes 1-5; second 
row, nodes 6-8, third row, nodes 9-12, fourth row, nodes 13-16, top row, nodes 17-18 (18 at 
base of terminal spike). INSET: successive leaves from lateral branch shown by arrow in habit 
diagram. (Silhouettes photographed by T. F. Beals). 


314 Rhodora [ Vor. 60 


artemisiifolia X trifida, and it is possible that the production 
of any fruits at all is atypical. Nevertheless, our natural hybrid, 
cited above, that was cultured in 1957 under particularly luxu- 
riant conditions in the greenhouse and permitted to develop into 
an unusually large speeimen was grown side by side with numer- 
ous pollinating plants of A. artemisiifolia, and by the latter half 
of September the hybrid did produce approximately twenty 
fruits—these having been stimulated to form, perhaps, by the 


lia. 2. Fruits of ragweeds. A. Ambrosia coronopifolia; B. A. artemisiifolia; C. A. artemisii- 


olia X trifida; D. A. trifida. 


pollen from the parent species. The fruits, the outer walls of 
which are involucral in. origin. and appeared superficially to 
be normal, represent good intermediates between the rather 
different fruits of the parents as shown in the scale drawings in 
figure 2. The fruits of A. artemisiifolia average usually 4-5 mm. 
in total length, with a narrowly constricted beak that makes up 
one-third to one-half the length. Those of A. trifida are much 
larger, approximately double in over-all length, 7-10 mm., with a 
more broadly based beak that comprises one-fifth to one-fourth 
of the total length. The fruits of the hybrid are variable but all 
of those which did develop on the plant were intermediate both 


1958] Wagner,—Hybrid Ragweed 315 


in size and shape between those of the parents. The only major 
deviation is that the hybrid fruits tend to be more deeply chan- 
nelled, a feature no doubt related to the collapse or failure of 
development of the achene within. Following after-ripening 
during the winter of 1957-58, an attempt was made to germinate 
some of the hybrid fruits, but without success. Both the pollen 
and the fruits (when formed) of this hybrid are probably ineffec- 
tive in its reproduction. 

Because of the great rarity and sporadic occurrence of this 
hybrid ragweed, and because of its probable inability to repro- 
duce or even survive more than one summer by any means, I am 
inclined to designate it by formula only, i.e., as Ambrosia arte- 
mistifolia X trifida, rather than by formal binomial nomenclature, 
as A. X helenae. There is considerably more justification, per- 
haps, for designating the ragweed hybrid, A. artemisiifolia X 
coronopifolia with a taxonomic binomial as A. X intergradiens 
(Wagner & Beals, op. cit.), because the latter hybrid is found to 
be frequent in many counties of Michigan (and probably else- 
where) and has the ability to form very large, perennial popula- 
tions through the years by way of root proliferations. The 
question, however, of when or why it is “useful or necessary" to 
designate an interspecific plant hybrid with a binomial is at 
present wholly arbitrary and personal. It would be desirable if 
there existed a generally acceptable and sound philosophy as a 
guide in this regard. 


LITERATURE CITED 


Jones, G. N. and G. D. Futter. 1955. Vascular plants of Illinois. 
Univ. Illinois Press, Urbana. 

Jones, K. L. 1943. Studies in Ambrosia: III. Pistillate Ambrosia 
elatior X A. trifida and its bearing on matroclinie sex inheritance. 
Bot. Gaz. 105: 226-231. 

KuMMER, ANNA P. 1951. Weed seedlings. Univ. of Chicago Press, 
Chicago. 

LawALRÉE, ANDRE, 1947. Les Ambrosia adventices en Europe occi- 
dentale. Bull. Jard. Bot. de l'Etat Bruxelles 18: 305-315. 

RouLEAv, Ernest. 1944. Notes taxonomiques sur la flore phanéro- 
gamique du Québec. —1. Naturaliste canadien 71: 265-272. 

Rower, W. W. 1893. Studies upon akenes and seedlings of the order 
Compositae. Bull. Torr. Bot. Club 20: 1-17 

RYDBERG, P. A. 1922. Ambrosiaceae, N, Am, Flora 33, part 1: 3-44. 


316 Rhodora | Vor. 60 


STEBBINS, G. L., Jr. 1945. The evtological analysis of species hybrids. 
II. Bot. Rev. 11: 463-486. 

Waaner, W. IL, Jn. and T. F. Bears. 1958. Perennial ragweeds 
(Ambrosia) in Michigan with the description of a new, intermediate 
taxon. RHODORA 60: 177-204. 

Wyvrig, R. B. 1915. A hybrid ragweed. Proe. Iowa Acad. Sei. 22: 
127-128, pl. 21. 


SOME INTERESTING RECORDS FROM EASTERN 
MASSACHUSETTS 


RICHARD J. EATON 


Middlesex and Norfolk Counties in eastern Massachusetts, par- 
ticularly within a radius of thirty miles from downtown Boston, 
have been intensively explored during the past century by several 
generations of energetic botanists. The flora of that area is, 
perhaps, as well represented by herbarium specimens, and the 
detailed distribution of its specific elements is as well known, as 
that of any comparable area in the New World. It is, therefore, 
à source of constant interest and surprise that novelties (from a 
local point of view) continue to crop up in some numbers. De- 
spite the accelerating contraction of suitable habitats for our 
native vegetation and the rapid conversion of rural areas to 
industrial, highway, and housing uses with all that that process 
implies, it is still profitable to rework familiar ground intensively 
in search of previously overlooked plants or recent arrivals. 

The specimens cited below were collected by the author unless 
otherwise stated, and have been placed in the herbarium of the 
New England Botanical Club. 


Isopres RIPARIA Engelm. Walden Pond, Concord, Middlesex County, 
Massachusetts, Sept. 22, 1957. Neither it nor its more northerly and 
hardly separable var. canadensis has been reported previously from the 
county. Even if considered as an aggregate, this species appears to be 
somewhat local in eastern New England: one record each from Cumber- 
land and York Counties, Maine; one from Suffolk, three from Norfolk, 
and two from Worcester Counties, Massachusetts; two from Providence 
County, Rhode Island; two from Windham County, Connecticut. 

IsoErES MACROSPORA Dur. Walden Pond, Concord, Massachusetts, G. 
HR. Proctor 3681-b, Aug. 29, 1949; same station, Sept. 22, 1957. New to 
Middlesex County, Massachusetts. Aside from three other stations in 
Massachusetts (Plymouth and Hampden Counties) this species appears to 
be confined to northern New England. Unaware of the Proctor collection, 


1958] Eaton,—Records from Eastern Massachusetts 317 
I stumbled on it last fall while making a periodic “low-water” visit to 
Walden. It seems odd that the numerous keen botanists who have ex- 
plored its shores and neighboring woods during the past hundred years 
should have previously overlooked two species of Isoëtes. The suspicion 
is inevitable that they are recent arrivals, particularly in view of the ever- 
inereasing hordes of bait fishermen and others who infest the pond. 

CYPERUS ERYTHRORHIZOS Muhl. Two recent collections from Concord: 
Great Meadows, J. W. Brainerd 2569, July, 1950; shore of Walden Pond, 
Sept. 22, 1957. Previously known in Middlesex County only from Winter 
Pond, Winchester. Until 1930 this species was considered rare in southern 
New England where it reaches its northeast limit of range at two stations 
in Massachusetts on the bank of the Merrimaek River in Essex County. 
It was known elsewhere in our area at two stations each in Norfolk and 
Bristol Counties, all in Massachusetts, and at a scattering of seven sta- 
tions in the Connecticut River valley from Hadley, Massachusetts, to its 
mouth at Lyme, Connecticut. Since 1930, in addition to the two Concord 
stations cited above, it has been discovered at five places in the Black- 
stone River valley from Auburn (near Worcester), Massachusetts, south- 
ward to Providence, Rhode Island; and at two places along the upper 
(southern) Nashua River in eastern Worcester County. It is difficult to 
accept the idea that this well-marked and easily recognizable sedge pre- 
viously had been overlooked in such a thoroughly botanized town as 
Concord.! 

More problematical is its status in Worcester County, Massachusetts, 
where active and competent botanists under the leadership of B. N. Gates 
and David Potter have been systematically collecting during the past 
twenty-five years. However, Joseph Jackson did intensive field work 
there (but without the benefit of motor transportation) in the 1890's 
while gathering material for his “Flora of Worcester County”. During 
the next two decades, in addition to continued field work by local botanists, 
Messrs. Fernald, Knowlton, Hunnewell, and other members of the New 
England Botanical Club, either on formal field excursions or otherwise, 
visited the county more or less regularly. In short, its flora was reason- 
ably well known to botanists of a preceding generation. Therefore, one 
must not dismiss the possibility, if not the probability, that C. erythrorhizos 
has invaded the Sudbury-Concord, Nashua, and Blackstone üver valleys 
in recent years. If so, the reasons and agencies responsible for its possibly 


1 The New England Botanical Club Herbarium contains specimens from Concord repre- 
senting more than 1050 species plus an additional 124 named varieties and forms, collected by 
forty persons beginning with H. D. Thoreau and E. S. Hoar in the 1850's. 

After these notes were sent to the printer an old undated specimen has been found in the 
recently acquired herbarium of Mary Rodman. The label is inadequately annotated 
“Bedford—C. W. J.” Evidently she received it from Charles W. Jenks, a contemporary 
amateur botanist of Bedtord who collected actively in central Middlesex County beginning 
about 1880. It is physically possible that Jenks could have collected the plant in the 
Bedford section of the Concord River meadows, but there are good reasons for disregarding 
this possibility. Internal evidence strongly suggests that the specimen did not originate 
with Jenks or from Bedford. 


318 Rhodor: [ Vor. 60 


sudden dispersal must remain obscure. One is tempted to correlate the 
postulated phenomenon with the remarkable increase in eastern Massa- 
chusetts of migrant waterfowl, in particular surface-feeding fresh water 
ducks, such as Black Duck, Wood Duck, Pintail, Blue- and Green-winged 
Teal? This correlation probably is coincidental. The recent increases, 
in fact, merely represent partial recoveries from the steady decline of 
waterfowl along the eastern flyway since early Colonia! times when their 
abundance appears to have been fantastic. Obviously, the opportunities 
for dispersal of the sedge by avian means in post-glacial times were far 
greater prior to 1700, for instance, than during the years 1930-1957. 

CAREX HIRSUTELLA  Mackenz. Open deciduous woods, Concord, 
Massachusetts, June 16, 1957. First record from Middlesex County. 
I find only two herbarium specimens from eastern New England: one 
from Mt. Desert, Maine, and one from Bristol County, southeastern 
Massachusetts. Otherwise, it appears to be confined to localities west of 
the Connecticut River from Vermont to Connecticut. Dr. F. J. Hermann 
concurs with my determination of the Concord specimen. 

CAREX LAXIFLORA Lam., var. SERRULATA F, J. Herm. | Lincoln, Middle- 
sex County, Massachusetts, June 22, 1955, wooded rocky slope over 
diorite outcropping. Obviously closely related to C. laxiflora, but unlike 
anything to be expected in eastern Massachusetts, I sent the specimen to 
Dr. Hermann for his opinion. He assigned it unhesitatingly to this ap- 
parently local and seldom collected variety. This appears to be its first 
recorded occurrence in New England. There are only five specimens in 
the Gray Herbarium: one each from New York, Michigan, and Washing- 
ton, D. C., and two from Tennessee. Dr. Hermann writes that he has 
seen a recent collection from an additional station in Michigan. Like the 
typical form, var. serrulata seems to prefer rich hardwoods. Incidentally, 
the Lincoln station, according to the detailed geological map of the region, 
lies on the same formation as that for Parietaria pensylvanica in Concord 
mentioned below. "Phe formation is mapped as a narrow, gradually ex- 
panding ribbon from Marlborough to Salem, Massachusetts. Except 
near the coast, it reaches the surface as outerops at but a few places; 
otherwise it is buried chiefly by the acid soils of glacial origin. The out- 
cropping in Concord is noteworthy for its rich flora. It is the site of a 
number of species not known to occur elsewhere in the town and which 
are relatively rare and local in eastern Massachusetts. To a much lesser 
extent, the same is true of the less extensive Lincoln exposure. 

Wo ria puncrata Griseb. Concord, Massachusetts, Oct. 16, 1957, on 
quiet water in an artificial wild-fowl impoundment adjacent to the Great 
Meadows impoundments of the United States Wild Life Refuge. The 
presence of this species was discovered by Carroll E. Wood, Jr., while 
searching for fruiting fronds of W. columbiana in a sample I had collected 
for him for that purpose. A collection subsequently taken from a de- 
tached colony of the now locally abundant W. columbiana? contained 

? See “Birds of Concord" by Ludlow Griscom (Cambridge, 1949), pp. 153-154, 181-188, ete. 


3 First discovered in eastern Massachusetts in 1938 and reported by the author in RHODORA, 
41: 42 


1958] Eaton,—Reecords from Eastern Massachusetts 319 


roughly five per cent W. punctata. The latter species seems to float 
slightly higher in the water than the other and is somewhat boat-shaped. 
By running a cupful through a fine-meshed kitchen strainer to remove 
contaminating Lemna, ete., I was able to obtain a pure sample of the mixed 
Wolfiia plants. Successive portions of the cleaned material were floated 
in a flat dish. By gently blowing across the surface, I found it possible 
to increase the percentage of W. punctata at the far edge to a marked 
degree, thus facilitating the preparation of a pure specimen of the latter 
species.‘ 

This appears to be its first recorded occurrence in New England. Its 
range is stated by Fernald in Gray's Manual, Sth Ed., as “Fla. to Tex., 
n., very locally, to e. Md., n.w. N. Y., s. Ont., Mich ...", ete. In con- 
nection with the possibility of the dissemination of aquatic vegetation by 
waterfowl, suitable ecological conditions at the place of deposition are 
essential for a successful introduction. In the case of the Lemnaceae, the 
sewage-polluted waters of the Sudburv-Coneord River and particularly 
the Great Meadows impoundments adjacent to it are notoriously favorable 
for their luxuriant growth.? Seemingly, the chances are excellent that a 
single frond of Wolffia, released from entanglement in some portion of the 
external anatomy of a migrant duck, for instance, onto the surface of one 
of these impoundments, should soon proliferate abundantly. It is con- 
sidered well-nigh impossible for any vegetal material, viable or otherwise, 
to remain in the digestive tract of a bird during the time required for a 
flight from Maryland to eastern Massachusetts. Also, it is questionable 
whether Wolffia ever can stick to a bird's feathers. It is conceivable that 
it can stick to a strand of weed fouled in a bird's foot or bill. "Very rarely 
I have seen a land bird with a claw or bill apparently fouled by extraneous 
material. One or two of my ornithological friends have observed similar 
occurrences. Wolffia adheres strongly by capillary attraction to moist 
surfaces. Such a means of long-distance transport seems possible. On 
the whole, I think it rather likely that both species of Wolffia are recent 
arrivals in Concord. 

SISYRINCHIUM ARENICOLA Bickn. Braintree, Norfolk County, Massa- 
chusetts, R. B. Channell and R. J. Eaton, July 5, 1957. The previously 
known occurrence of this southern species in northeastern North America 
is indicated by specimens in the Club and Gray Herbarium from Wood- 
bury, Connecticut, Block Island, Rhode Island, Marthas Vineyard, 
Nantucket, and Provincetown in Massachusetts, and thence (disruptedly) 
from southwestern Nova Scotia. This familiar distributional pattern is 
characteristic of a number of southern plants which are assumed to have 
migrated in the late Pleistocene northeastward along the exposed con- 
tinental shelf, and to have become isolated at scattered marginal stations 
by subsequent rise in the ocean level. Hence, the occurrence of sS. 


4 For methods of preserving herbarium specimens of Wolffia see “Wolffia in Canada" by 
W. G. Dore in The Canadian Field-Naturalist, 71: 10-106. 

? See author's paper “Lemna minor as an aggressive weed in the Sudbury River". RHODORA, 
49: 165-171. 


320 Rhodora | Vor. 60 


arenicola on the mainland of Massachusetts in Norfo!k County is some- 
what surprising. 

PARIETARLA PENSYLVANICA Muhl. Deciduous woods, base of dioritic 
outerop,® Concord, Massachusetts, Aug. 1, 1957. Previously unrecorded 
from Middlesex County. Except for three old records from the southerly 
outskirts of metropolitan Boston (one of which is definitely annotated 
"waste ground") its distribution in Massachusetts as reflected by the 
many specimens in the New England Botanical Club Herbarium is con- 
fined to the Connecticut River valley and westward, where rich woods 
species are notably numerous and abundant. The plant at Concord has 
every appearance of being indigenous. However, reference is made to 
Ruopona 1: 168-172 wherein P. pensylvanica is listed as one of the species 
said to have been unsuccessfully introduced at Concord by Minot Pratt 
(who died in 1878). Pratt, contrary to his usual practice, made no men- 
tion of the introduction in his “List of Plants of Concord" (ms. in the 
Concord Publie Library). 

BARTONIA PANICULATA (Michx.) Muhl. Shore of Bateman's Pond, 
Concord, Massachusetts, July 25, 1957. First record for Middiesex 
County. Previously known New England distribution: Cumberland and 
York Counties, Maine, one station each; Norfolk County, Massachusetts, 
three stations in the Blue Hills Reservation; thence westward near the 
coast in southeastern Massachusetts, Rhode Island and Connecticut. 

CRATAEGUS CRUS-GALLI L. Concord, Massachusetts, July 25, 1957. 
Second record for the county, the first being Middlesex Fells, N. T. 
Kidder, June 8, 1928; otherwise not known in eastern Massachusetts, 
north of Bristol County. This species is one of the very few New England 
members of the genus which ean be readily identified in the field. At the 
Concord station were three large and apparently old individuals and sev- 
eral smaller ones at the edge of extensive rocky open woodland in a long 
abandoned pasture. 

VERNONIA MISSOURICA Raf, Clam Shell Bluff, Concord, Massachusetts, 
Aug. 21, 1957. Adventive from the West. Probably the first record for 
New England. It is included here as a previously overlooked addition to 
the long list of adventives at this station reported by the author 
in Ruopora 38:64-67. It has presumably persisted for nearly thirty 
years, despite the fact that all but narrow edges of the field recently has 
been put back into cultivation. 

HIERACIUM PILOSELLA L. var. NIVEUM Muell. Arg. Old. sterile. field, 
Concord, Massachusetts, July 30, 1957, where it occurs as a dense clone 
several meters in diameter. Naturalized from Europe. Its occurrence 
in the Gray's Manual range is stated by Fernald as "Fields, e. Maine”. 
Var. niveum strikingly differs from the frequently collected and wide- 
ranging typical form in that its leaves are permanently and conspicuously 
white pannose beneath. Furthermore, it appears to be far less aggressive. 
— LINCOLN, MASS. 


5 The pH of a finely powdered mixed sample was determined as 6.2. 


1958] Gates,—Cabomba Caroliniana 321 


CABOMBA CAROLINIANA GROWS IN WORCESTER 
COUNTY, MASSACHUSETTS 


Burton N. GATES 


The distribution of Cabomba caroliniana A. Gray in Massa- 
chusetts is gradually unfolding. An important recent contribu- 
tion is that of Stuart K. Harris! who collected it in Fosters Pond, 
Essex County, 1957. In the next County to the south, Suffolk, 
Dr. Harris also reported it in abundance in Muddy River, The 
Fenway, Boston. South of Boston, the present writer has seen a 
specimen from the sandy shore of Nippenicket Pond, Bridge- 
water, Plymouth County (Frank C. Seymour, 4426, 1935; a 
sterile specimen in the New England Botanical Club Herbarium). 

The range of Cabomba extends westward into Worcester 
County. The specimens documenting this distribution are filed 
in the Hadwen Herbarium of Worcester County Plants, Clark 
University, Worcester. The earliest collection was from Ux- 
bridge, in the Mumford River (Walter H. Hodge, 2825, 1933). 
Again in Uxbridge, it was found in great abundance; the long 
streamers,” having been washed up by a hurricane, were floating 
along the shore of Ironstone Reservoir. A random specimen 
measured 85+ inches long (F. C. Seymour and B. N. Gates 32316, 
1954). Nine years previous, it had been collected as an occa- 
sional plant in Ironstone Brook at Ironstone Road, which is fed 
by Ironstone Reservoir (B. N. Gates 23914, 1946). In the 
southeast town in Worcester County, Blackstone, in the Black- 
stone River, two or three feet deep, Cabomba covered the bed of 
the river (Malumphy 24893, 1943). 

Distribution in Worcester County extends west considerably 
beyond the geographical center of the State. On a field trip of 
the New England Botanical Club it was first collected floating 
along the north-east shore of Quaboag Pond near the inlet of 
Seven Mile River (David Potter 24913, 1946). At this station, it 
has been observed nearly every year since and was collected in 
May 1955 (Winifred C. Gates 31976). That there is no associa- 
tion by water-courses between this East Brookfield station and 

! Ruopora 60: 116. 1958. 

‘A profusion of amazingly long streamers ot Myriophyllum humile forma capillaceum tangled 


with Cabomba; the collected specimen measured in excess of 11.5 feet long (F. C. Seymour and 
B. N. Gates 31995). 


322 Rhodora | Vow. 60 


the two towns, Uxbridge and Blackstone, is quite evident; East 
Brookfield is in the water-shed which empties into the Thames 
River in Connecticut, while the two other towns are in the 
Blackstone River water-shed, draining south through Rhode 
Island. 

Westward from Worcester County, Fassett,’ without a specific 
citation, lists an occurrence in “western Massachusetts, where 
perhaps introduced", is interpretable as most any locality west of 
the Connecticut River. It is quite probable, however, that his 
citation refers to the collection in 1930 by Wayne E. Manning? 
in Hatfield, Hampshire County. He found it very abundant in 
South Pond, once a very old ox-bow of the Connectieut River. 
A local florist and fisherman, Harold Keys, is quoted by Manning 
as having known it in profusion in this pond for at least a decade 
previous and as having suggested that it appeared to be 
indigenous. 

To round out the statement of occurrence of Cabomba in 
southern New England, it may be helpful to cite briefly two col- 
lections in Connecticut, which have been seen by the writer. 
Ultimately, it might develop that they have some relationship 
to the Massachusetts station in the Connecticut River Valley 
water-shed. In Rogers Pond, Saybrook, Middlesex County, it 
was reported to be very abundant (E. H. Hames 11854a, 1937; a 
specimen in flower at the Gray Herbarium). In a pond at the 
south-west base of Chestnut Hill, Trumbull, Fairfax County, it 
has been long established, according to the property owner. 
(E. H. Eames 11874, 1937; a specimen in flower in the Gray 
Herbarium). 

Sterile Cabomba has a reasonable resemblance to several other 
aquatics found in our New England waters. For this reason 
and because in the writer’s experience it is seldom found in flower 
or fruit, at least in Massachusetts, it may be readily overlooked 
or mistaken for one of several plants of ponds and streams. 
Sterile specimens of Cabomba have the general appearance of 
Ceratophyllum demersum L., Megalodonta Beckii (Torr.) Greene, 
Ranunculus flabellaris Raf. and to some forms of Myriophyllum. 
Distinguishing characters of the stem and leaves of these are very 
helpfully pictured by W. C. Muenscher, in comparison with the 


! Manual of Aquatic Plants, 1940. 
! Ruopora 39: 187, 


1958] Proetor,—Further New Records of Myrtaceae 323 


stem and leaves of Cabomba, on a plate in his Aquatic Plants 
of United States”. Observe that Cabomba has a well developed 
petiole without stipules. 

Conclusions are few, beyond the fact that Cabomba is locally 
widespread in Massachusetts. The earliest collection known in 
Massachusetts was in 1930, in Hatfield, Hampshire County, fol- 
lowed in Worcester County, in Uxbridge, in 1933. In the writer's 
experience, supported by the collections of others, it is locally 
plentiful and well established. Regrettably, the writer has yet 
to see it in flower or fruit in the area (although the two Connecti- 
eut collections were in flower). Occurrence in Massachusetts 
constitutes an extension considerably north of the normal range 
given in Gray's Manual, ed. 8, as Virginia, or, by Fassett, as 
New Jersey. Each of these authors would account for the New 
England extension as “naturalized” or “perhaps introduced" 
through some unexplained means. Its scattered but established 
distribution anticipates that further collections should show a 
more unified occurrence of Cabomba in southern New England.— 
BIOLOGY DEPARTMENT, CLARK UNIVERSITY, WORCESTER, MASS. 


FURTHER NEW RECORDS OF MYRTACEAE 
FROM JAMAICA 


GEORGE R. Pnocron 


Recent botanical exploration by the writer in Jamaica has re- 
vealed an interesting series of new plant records. A previous 
number of Ruopona (50: 303-306. 1957) presented a few notes on 
Eugenia and proposed one new species. The present paper con- 
tributes more new Jamaican records in the Myrtaceae. "The 
writer is grateful to Dr. Lily M. Perry for her assistance with the 
Latin descriptions. 

Calyptranthes clarendonensis Proctor, sp. nov. Frutex cirea 2.5 m. 
altus; ramulis novellis fere teretibus, dense brunneo-tomentosis. Folia 
sessilia, rigide coriacea, opaca, non pellucido-punctata, supra glabra, 
subtus decidue brunneo-tomentosa, 2.5-5 em. longa, 1.5-4.5 em. lata, late 
ovata, apice obtusa, basi valde cordato-subamplexicaule, nervo medio 
supra basin versus impresso, subtus prominente, venis secundariis promi- 
nulis. Inflorescentiae 2-4, subterminales, apice 2-3-florae, floribus sessili- 
bus. Alabastra ovoidea, apiculata, 5-6 mm. longa, dense brunneo- 


5 [thaca, N. Y., 1944, plate 102a, page 232, 


324 Rhodora [ Vor. 60 


tomentosa. Flores aperti non visi. Baccae  globosae (interdum 
irregulariter globosae), brunneo-tomentosae, 9-14 mm. diametro; semini- 
bus 2-13, 6-9 mm. longis. 

Shrub c. 2.5 m. tall; youngest branchlets nearly terete, densely brown- 
tomentose. Leaves sessile, rigidly coriaceous, opaque and without pellu- 
cid dots, glabrous above, deciduously brown-tomentose beneath, 2.5-5 
em. long, 1.5-4.5 em. wide, broadly ovate, blunt at apex and rather deeply 
cordate-clasping at base, midrib impressed toward the base above, promi- 
nent beneath, secondary venation prominulous on both sides. In- 
florescences 2-4, subterminal (i.e., accompanied and overtopped by a pair 
of leafy shoots which appear simultaneously from the same place); pe- 
duncles 2-5 em. long, somewhat compressed or 2-edged, brown-tomentose, 
'ach bearing 2 or 3 sessile flowers at the apex. Buds ovoid-apiculate, 5-6 
mm. long, densely brown-tomentose. Open flowers not seen. Berries 
globose (sometimes irregularly), brown-tomentose, 9-14 mm. in diameter, 
2-13-seeded, the seeds 6-9 mm. long. 

TYPE: Parish of Clarendon, Peckham Woods, elev. e. 2500 ft., on roeky 
limestone hilltop, Proctor 11399, colleeted Dee. 23, 1955. Holotype at 
the Institute of Jamaica; the type material is in fruit. 

ADDITIONAL SPECIMENS: from the same locality, Webster & Proctor 5392, 
with young fruits (Herb. Arnold Arboretum); Proctor 9760, January 7, 
1955, with buds and fruits. 

The present new species was for a time doubtfully identified 
with C. maxonii, but clearly differs from that species in its terete, 
tomentose branchlets; larger and more deeply cordate leaves; 
longer peduncles bearing but 2 or 3 sessile flowers (instead of ‘3 
heads of (to 12) sessile flowers, the 2 lateral heads more or less 
stalked”); and much larger buds. The fruits of C. maronii 
evidently are not known; those of C. clarendonensis tend to have 
more seeds than is usual in this genus. 

From C. cardiophylla Urb. of eastern Cuba, with which C. 
clarendonensis seems to show a near relationship, the latter differs 
by the leaves entirely lacking the “densely impressed-punctate”’ 
character of the upper surface and by differences in the nature of 
the pubescence. The flowers and fruits of C. cardiophylla were 
unknown to Urban, but more recently-collected material (Alain 
3352), with buds, suggests that these structures are smaller in the 
Cuban species. 

Calyptranthes ekmanii Urban, Ark. Bot. 224 (10): 32. 1929. Sterile 
material apparently conspecific with this small tree of southwestern Haiti 
has been collected in moist elfin woodland on the east slope of the John 


Crow Mountains, Parish of Portland, Proctor 9820. This is a new record 
for Jamaica, 


1958] Proctor, —Further New Records of Myrtaceae 325 


Myrcia skeldingi Proctor, sp. nov. Arbuscula circa 5 m. alta; ramulis, 
petiolis, nervo medio foliorum subtus, et inflorescentiae ramis superioribus 
sparsim strigillosis, deinde glabratis. Folia coriacea et sine punctis pellu- 
cidis, subsessilia (petiolis crassis, 1-2 mm. longis), 5.5-10 em. longa, 3.5-6 
em. lata, late elliptica vel ovato-elliptiea, apice plerumque rotundata vel 
obtusa, basi subeordata vel rotundata, nervo medio supra impresso, subtus 
prominente, venulis crebris et tenuibus reticulatis utrinque sed praecipue 
subtus prominulis. Paniculae terminales multiflorae, plerumque 7-14 em. 
longae lataeque, peduneulo 0.5-1 cm. longo, ramis ultimis plerumque 
dichasiam 3-5-floram terminalem ferentibus. Tubus calycis 1.5 mm. 
longus, glaber, paulum supra ovarium productus, 4-lobatis, lobis circa 0.75 
mm. longis; petalis albis, glabris, ca. 1 mm. longis; antheris bilocularibus; 
stylo ca. 3.5 mm. longo, stigmate peltato. Baccae globosae, glabrae, ca. 6 
mm. diametro, maturae rubrae, seminibus 1-3, (3)-4 mm. longis. 

Small tree c. 5 m. tall; youngest branchlets, petioles, midribs of leaves 
beneath, and upper infloreseence-branches sparsely strigillose, all parts 
becoming glabrate with age. Leaves coriaceous and without pellucid dots, 
subsessile (petioles thick, 1-2 mm. long) 5.5-10 em. long, 3.5-6 em. wide, 
broadly elliptie or ovate-elliptie, the blades mostly rounded or blunt at the 
apex, with base subcordate or rounded; midrib impressed above, prominent 
beneath, the finely-reticulate venation prominulous on both sides but 
especially beneath. Panicles terminal, many-flowered, mostly 7-14 em. 
long and broad, the basal stalk (peduncle) 0.5-1 em. long, the ultimate 
branches mostly ending in 3-5-flowered dichasia. Calyx with tube 1.5 
mm. long, glabrous, slightly prolonged above the ovary, 4-lobed, the lobes 
c. 0.75 mm. long; petals white, glabrous, c. 1 mm. long; anthers 2-celled ; 
style c. 3.5 mm. long, with peltate stigma. Berries globose, c. 6 mm. in 
diameter and deep red when ripe, with 1-3 seeds, these (3)-4 mm. long. 

TYPE: Parish of Clarendon, Mason River Savanna, 2.75-3 miles north- 
west of Kellits, elev. e. 2300 ft., in thickets along stream, Proctor 16478, 
collected July 9, 1957. Holotype at the Institute of Jamaica; the type 
material has flowers and young fruits. 

ADDITIONAL SPECIMEN: from the same locality, Proctor 16734, collected 
Oct. 14, 1957, with ripe fruits. 


Myrcia skeldingi is easily distinguished from other J amaican 
congeners (and most species elsewhere) by its subsessile, more or 
less subeordate, blunt-tipped leaves and 4-lobed calyx. In the 
latter character it is an anomalous member of its genus. This 
species shows a rather close resemblance to a group of Brazilian 
forms, especially M. uberavensis Berg (=M. cardiophylla 
Reicht. ?), but the latter clearly differ in having 5-parted flowers 
and in other individual details. 

Named for Prof. A. D. Skelding of the Department of Botany, 
University College of the West Indies, who first pointed out to the 


326 Rhodora [ Vor. 60 


writer the botanical richness of the Mason River Savanna area 
where this species is apparently endemic. 

Eugenia jeremiensis Urb. & Ekm., Ark. Bot. 244 (4): 29. 1931. Ja- 
MAICA: Parish of Hanover, interior summit slopes of Dolphin Head, on 
moist wooded limestone hillside, Proctor 10032 and 10416. 

The specimens cited above were originally believed by the 
writer to represent an undescribed species, but subsequent com- 
parison with type material of E. jeremiensis (from southwestern 
Haiti) strongly suggests that they are conspecific. Ekman’s 
specimens of the latter species in the Institute of Jamaica her- 
barium are, however, sterile, and the Jamaican plants may yet 
prove to be different. In any case, this material represents : 
new record for Jamaica. It can be described as follows: 

Shrub or small tree reaching at least 5 m. in height; branchlets 
terete or slightly compressed near the end, glabrous. Leaves 
coriaceous, glabrous, 10-12 em. long by 7-9 em. broad, ovate- 
orbicular with rounded apex, the base of the blade broadly 
cuneate, dark green above and paler beneath with numerous 
minute faintly pellucid punctate dots; midrib slightly impressed 
near the base on the upper side, but flat or slightly raised toward 
the apex, very prominent beneath, especially near the base, with 
side-veins and venules prominulous on both sides; petioles thick, 
furrowed above, 2-3 mm. long. Flowers paired in the upper 
leaf-axils; pedicels 2.5 em. long, stout (1-1.5 mm. in diameter); 
bracteoles united at the base, broadly deltoid-ovate, obtuse, 1 
mm. long; sepals apparently roundish, unequal, the longer 1.4 em. 
long, the shorter 1 em. long (seen in bud only). Expanded 
flowers and fruits not seen. 

The flowers of this species, when open, must be larger than 
those of any other indigenous Éugenia.—1iNsTITUTE OF JAMAICA, 
KINGSTON, JAMAICA, W.I. 


1958] Wilson,—Typifieation of Genus Forestiera 327 


TYPIFICATION OF THE GENUS FORESTIERA 
(OLEACEAE)! 


In preparing the treatment of the genera of the Oleaceae of the 
southeastern United States, I found it necessary to determine the 
type of the genus Forestiera. Two different species have already 
been chosen as the type, and the problem was to discover which 
one was correct. Forestiera acuminata (Michx.) Poir. was desig- 
nated by Britton and Brown (Ill. Flora ed. 2. 2: 278. 1913) and 
by A. Rehder (Bibl. Cult. Trees & Shrubs p. 576. 1949) as the 
type of the genus while M. C. Johnston (Synopsis of the United 
States species of Forestiera (Oleaceae). Southwestern Nat. 2: 
141. 1957 [1958]) maintained that the type species is Adelia 
porulosa Michx. After reviewing the literature, however, I have 
reached still another conclusion. 

The genus Adelia was described by Patrick Browne in 1756 
(Civ. Nat. Hist. Jamaica p. 361), but he listed only the poly- 
nomial *ADELIA I. Foliis obovatis, oppositis; spicillis alaribus 
[sic]; cortica cinereo," and made no binomial combination. He 
did, however, include a short description and an illustration of 
the male plant (Tab. 36, fig. 3) which he said was “common in 
the low gravelly hills eastward of Kingston.” 

Michaux (Flora Boreali-Americana 2: 223. 1803) adopted the 
name Adelia (“ADELIA. Brown.") and described three species, 
making the proper binomial combinations. These species were 
Adelia porulosa ("*HAB. in maritimus Floridae"), A. ligustrina 
(*HAB. in fruticetis Illinoensibus, Tennasée, &c"), and A. 
acuminata (*HAB. ad ripas fluviorum Carolinae et Georgiae). 
Browne's plant was not named by Michaux. 

Willdenow (Sp. Pl. ed. 4. 4: 711. 1806) proposed the name 
Borya for the genus, and listed “Adelia Mich. amer. 2. p. 223” in 
synonymy. He included 4 species in his treatment: Borya 
cassinoides Willd., based on Browne's description and also a 
Richard specimen (‘Habitat in Antillis"); Borya porulosa Willd., 
based on Adelia porulosa Michx. (“Habitat in maritimis Flori- 
dae"); Borya ligustrina, and Borya acuminata. Willdenow, 
therefore, included Michaux's three species in his treatment, but, 

! Continuing a series of miscellaneous notes and papers on the flora of the southeastern 


United States made possible through the interest and support of George R. Cooley and a grant 
from the National Science Foundation. 


328 Rhodora [ Vor. 60 


in addition, described and named Browne's plant upon which the 
genus Adelia was based. The name Borya Willd., however, was a 
later homonym of Borya Labill. (Liliaceae). 

Forestiera was the name proposed for this genus in 1810 by 
Poiret (Encycl. Méth. Suppl. 1: 132. 1810) who cited both Adelia 
Michx. and Borya Willd. in the synonymy. Among the four 
species listed by Poiret (op. cit. 2: 663-665. 1811 [1812]) are 
Forestiera cassinoides, based on Borya cassinoides Willd. and 
Browne's description and figure (“Cette plante croit aux Antilles. 
(Herb. | Richard.)") and  Forestiera porulosa, based on Borya 
porulosa Willd. and Adelia porulosa Michx. (“Cette plante croît 
dans la Floride, sur les cótes de la mer.’’). 

It is clear, then, that Patrick Browne's plant bears the specifie 
epithet cassinoides, (not porulosa). Moreover, since this is the 
species upon which P. Browne's Adelia is based (and therefore 
also Borya Willd. and Forestiera Poir.) it should be considered as 
the type of the genus. Since Adelia L. (Euphorbiaceae) has been 
conserved over Adelia Browne, the correct name for the genus is 
Forestiera Poir., but the type remains the same. 

The type species of the genus Forestiera is Borya cassinoides 
Willd. (= Adelia cassinoides (Willd.) O. Ktze. =Forestiera cas- 
sinoides (Willd.) Poir.).—KENNETH A. WILSON, GRAY HERBARIUM 
AND ARNOLD ARBORETUM. 


A NEW RHODODENDRON STATION IN MAINE.—As a 
result of an intensive botanical and horticultural study of native 
Rhododendron maximum colonies in Maine and New Hampshire, 
one hitherto unreported stand has come to our attention. Pro- 
fessor Elwyn Meader of the Department of Horticulture, Uni- 
versity of New Hampshire learned of it first and called it to the 
attention of the senior author. 

A visit was made by the authors and notes taken on November 
4, 1957. This is a very small colony consisting of rather uniform 
plants standing about 215 feet high and covering an area about 10 
feet long and 6 feet wide. It is situated a short distance in from 
the north shore of Horn Pond, Acton, York County, Maine, and is 
not more than !4 mile from the Maine-New Hampshire boundary. 
The environment is a mixed forest on a gentle south-facing slope. 
Deciduous species predominate but some large trees of Pinus 


1958] Blum and Wilce,— Type of Vaucheria compacta 329 


Strobus and Tsuga canadensis occur not far away. The Rhodo- 
dendrons are obviously young and it might be thought, therefore, 
that the colony is new and was established there recently either 
with the help of man or by means of natural migration. How- 
ever, there is no information that the colony was planted nor is it 
likely that an attractive ornamental species would be planted in 
this kind of situation. The nearest known natural colony is the 
well known one in Sanford which is more than 11 miles away. It 
seems improbable that R. maximum with its particularly relic 
character in New England traversed this distance in recent years. 

It may be more reasonable to conclude that there was formerly 
a larger colony in the area. Lumbering which is known to us as 
having been disastrous to other Rhododendron stands was 
formerly carried on at the Acton site and probably resulted in the 
almost complete destruction of the Rhododendrons. For- 
tunately, a few have survived. Now it will be interesting to see 
if it will regain its former abundance in future years.— RADCLIFFE 
Pike AND A. R. HODGDON, DEPARTMENT OF HORTICULTURE AND 
DEPARTMENT OF BOTANY, UNIVERSITY OF NEW HAMPSHIRE, 
DURHAM. 


THE TYPE or VAUCHERIA COMPACTA Var. KOKSOAKENSIS.—In describ- 
ing this variety on page 286 of Rhodora, Volume 60, we inadvertently 
neglected to cite the type specimen. The following should have appeared 
on that page. type: R. C. Wilce 830, collected on the eastern shore 
near the mouth of the Koksoak River, Québec, Canada, Sept. 1, 1955, in 
the herbarium of the New York Botanical Garden.—JOHN L. BLUM AND 
ROBERT T. WILCE. 


Volume 60, No. 719, including pages 289-308, was published 15 December, 
1958. 


330 


Rhodora | VoL. 60 


ERRATA 


Cover, No. 710, line 3; for Mid-arrowed, read Mid-arrowhead. 


Cover, No. 710, line 7 


; for Glen J. Winterringer read Glen S. Winterringer. 


Page. 34, line 7; for repens, read reptans. 
1 7 


Page 37, line 15; for 
Page 41, line 29; for 


repens, read reptans. 
listed either, read listed either because. 


Page 48, line 12; for Colorado, read southern Alberta. 


Page 58, line 4; for F 


‘asset, read Fassett. 


Cover, No. 711, line 9; for 76, read 86. 


Page 90, line 32; for 


Page 272, line 7; for 


alternis supra. 
Page 282, line 11; fo: 
Page 282, line 16; foi 
Page 282, line 27; foi 


constancy, read consistency. 
oppositis supra, alternis infra; read oppositis infra, 


" mm., read m. 
' 10-14 cm., read 10-14 mm. 
hirsute, read hispid. 


1958] Index to Volume 60 331 


INDEX TO VOLUME 60 


New scientific names and combinations are printed in bold face type 


Abies balsamea 248 

Acalypha rhomboidea 68 

Acer platanoides 65; saccharinum 
65; saccharum 65 

Achillea lanulosa 7, 8, 9; mille- 
folium 7, 8, 9, 66; Ptarmica 127 

Acnida altissima 301, 304; var. 
prostrata 301; var. subnuda 301, 
305; cannabina 301, 304; tamari- 
scina 301, 305 

Adelia acuminata 327; cassinoldes 
328; ligustrina 327; porulosa 327, 
328 

Adenoplea lindleyana 21 

Agropyron repens var. subulatum 
67 

Agrostis alba 65; canina f. mutica 
65; perennans 67; tenuis 65 

Ahles, H. A., Bell, C. R., and Rad- 
ford, A. E.. Species New to the 
Flora of North or South Caro- 
lina 10 

Ahmadjian, V.. The Fruticose and 
Foliose Lichens of Worcester 
County. Massachusetts 74 

Ajuga reptans 23 

Albino Form of Dipsacus sylves- 
iris, An 174 

Alectoria nidulifera 77 

Aletes acaulis 265. 266, 270; mac- 
dougalii 265, 270; humilis 265 

Alicabon barbadense 164 

Alliaria officinalis 15 

Alium ampeloprasum 12; 
atroviolaceum 12 

Alnus serrulata 206, 207 

Alsine rossi 45 

Alsinopsis macrantha 49 

Amaranthus albus 301, 302; ascend- 
ens 301, 305; caudatus 301. 305; 
cruentus 301, 305; deflexus 301, 
305; graecizans 14, 301, 305; hy- 
bridus 301. 305; f. hypochondri- 
acus 301. 305; lividus 301, 305; 
Palmeri 301. 305; Powelli 301. 
305; pumilus 301 ; retroflexus 301. 
304; spinosus 301, 305; viridis 
301, 305 


var. 


Ambrosia artemisiifolia 178-203, 
310-312. 314; f. villosa 194; var. 
elatior 68; bidentata 126. 201; 
coronopifolia 177-203; elatior 
178, 195, 201; media 195; psi- 
lostachya 177, 181, 183, 185, 188. 
190. 200—203. 311; var. coronopi- 
folia 126, 181; trifida 178, 180, 
181, 182, 195, 201, 310, 311, 312, 
314; artemisiifolia X coronopi- 
folia 315; A. x helenae 200, 315; 
x Ambrosia Helenae 310; ar- 
temisiifolia X trifida 309-315; 
bidentata X trifida 310; Ambro- 
sia X intergradiens 183. 186. 189- 
197. 200-203, 315; Plate 1234 

America, Arenaria rossii and Some 
of Its Relatives in 44 

Ampelamus albidus 21, 30 

Andropogon campyloracheus 10; 
gerardi 144; scoparius var. fre- 
quens 67 

Anaptychia hypoleuca 82 

Anastrophyllum saxicola 219, 220 

Antennaria aprica 306; campestris 
306; canadensis 306; fallax 306; 
microphylla 306; munda 306; 
neglecta 68, 306; neodioica 68, 
306; petaloidea 306; plantagini- 
folia 68, 306 

Anthelia juratzkana 217, 219. 223 

Anthemis arvensis var. agrestis 
127; var. arvensis 127; mixta 
127; tinctoria 127 

Anthoxanthum odoratum 65 

Anthriscus scandicinus 20 

Apios americana 70 

Apocynum androsaemifolium 70 

Aquilegia formosa var. flavescens 
86 

Arabis missouriensis 206 

Arceuthobium pusillum 298, 302 

Arctium minus 66 

Arenaria elegans 45, 47; filiorum 
49, 50, 51, 52; macrantha 44, 45. 
49. 50, 52; nuttallii 53; ssp. fragi- 
lis 53; ssp. gracilis 53; ssp. gre- 
garia 53; ssp. nuttallii 53; rosei 


332 Rhodora 


53; rossii 44—46. 49, 51; ssp. co- 
lumbiana 48, 50; var. apetala 
48, 50. 52; ssp. elegans 47. 51; 
ssp. rossii 45, 51; rossii var. ape- 
tala 49; var. columbiana 48; 
Rossii 45; rubella 50, 52; stricta 
45, 52; ssp. dawsonensis 53; ssp. 
macra 53; ssp. stricta 52; ssp. 
texana 52 

Arenaria rossii and Some of Its 
Relatives in America 44 

Aristida desmantha 42; lanosa 206. 
208; stricta 291 

Aristolochia Clematitis 298, 305; 
durior 298, 305; Serpentaria 298, 
303 

Arrhenatherum elatius f. biaristat- 
um 65 

Artemisia albula 127; dracuncu- 
loides 116; Parryi 266; vulgaris 
25 

Asarum canadense 298, 303; var. 
acuminatum 298, 303; var. re- 
flexum 298, 303 

Ascelepias syriaca 64, 68; verticil- 
lata 206 

Ascyrum hypericoides var. oblongi- 
folium 206 

Asparagus officinalis 65 

Asplenium cryptolepis 244 

Aster cordifolius 70, 72; ericoides 
68; infirmus 61; lateriflorus 68: 
var. pendulus 68; linearifolius 
68; novae-anghae 61, 66; novi- 
belgii 70; pilosus 143; puniceus 
var. compactus 70; undulatus 
68; vimineus 70 

Asterella ludwigii 211, 216—218. 224. 
234; saccata 211, 230. 231. 233. 
249. 250; tenella 217 

Athalamia hyalina 211. 230. 232. 
234. 247. 249. 250 

Athyrium Filix-femina var. Mich- 
auxili 69; f. elatius 69; f. laurenti- 
anum 69; thelypterioides f. 
acrostichoides 69 

Atriplex arenaria 300, 304; glabriu- 
scula 300, 304; hortensis 300. 305; 
patula 300, 304; var. hastata 300. 
304; var. littoralis 300. 304; rosea 
300, 305 

Axyris amaranthoides 300, 305 


| VoL. 60 


Baeomyces roseus 76 

Barbarea vulgaris var. arcuata 65 

Bartholomew’s Cobble (Review) 
32 

Bartonia paniculata 206, 319 

Bassia hirsuta 300, 305 

Bazzanin denudata 224 

Beals, T. F. and Wagner, W. H. Jr.. 
Perennial Ragweeds (Ambrosia) 
in Michigan. with the Deserip- 
tion of a New Intermediate 
Taxon 177 

Bean, R. C.. A. F. and Eaton, R. J., 
Twelfth Report of the Commit- 
tee on Plant Distribution 297 

Bell, C. R.. Radford. A. E., and 
Ahles, H. E., Species New to the 
Flora of North or South Carolina 
10 

Berberis vulgaris 65 

Bernard, Fr. J-P. and Lóve. D. 
Rumex stenophyllus in North 
America 54 

Beta vulgaris 300, 305 

Betula alba 65; lenta 71; populi- 
folia 67 

Bidens connata 58; frondosa 58. 
68; pilosa 25; var. radiata 25, 
30 

Blum. J. L. and Wilee, R. T. 
Description, Distribution and 
Ecology of Three Species of 
Vaucheria Previously Unknown 
from North America 283; Type 
of Vaucheria Compacta var. 
koksoakensis, The 329 

Boehmeria cylindrica 13, 206, 295. 
297. 302, 303; var. Drummondi- 
ana 297. 303; drummondiana 295 

Borya acuminata 327; cassinoides 
327. 328; ligustrina 327; porulosa 
327. 328 

Bothriochloa intermedia 91. 93, 95; 
ischaemum 91, 93, 94, 05; pertusa 
91. 93 

Brachiaria platyphylla 11 

Bredemevera colletioides 104 

Brodiaea uniflora 13 

Bromus brizaeformis 296; com- 
mutatus 65; marginatus 264 

Buchloé dactyloides in Illinois 259, 
260 

Buddleia lindleyana 21 


1958] Index to Volume 60 333 


Cabomba caroliniana 116, 321 

Cabomba caroliniana Grows in 
Worcester County, Massachu- 
setts 321; in Essex County, Mas- 
sachusetts 116 

Callirhoe involucrata 41. 43 

Callitriche terrestris 18, 30 

Calothrix scopulorum 284 

Calyptranthes cardiophylla 324; 
clarendonensis 323, 324; ekman- 
nii 324; maxonii 324 

Candelaria concolor 81; var. effusa 
81 

Cannabis sativa 297, 304 

Capsella Bursa-pastoris 65 

Carex annectens 67; cephalophora 
70; eleocharis 115; hirsutella 
318; laxiflora 318; var. serrulata 
318; lurida 206; normalis 70; 
pallescens var. neogaea 70; pen- 
svlvanica 70; var. digyna 115; 
picta 175; praegracilis 115; sco- 
paria 70; Shortiana 174; spicata 
65; squarrosa 174; stenophylla 
115; var. enervis 115; Swanii 70; 
tenera 70; typhina 174; virescens 
206; x Deamii 174 

x Carex Deamii in Missouri. 174 

Carolinas. Striga asiatica and 
Chrysoma pauciflosculosa in the 
289 

Carphephorus bellidifolius 121; 
carnosus 120, 121; corumbosus 
118, 121; pseudo-liatris 118, 121; 
tomentosus 121 

Carya glabra 67; ovalis 206; texana 
206; tomentosa 67. 206 

Castanea dentata 71 

Catalpa bignonioides 65 

Celarier, R. P. and Mehra. K. L.. 
Determination of Polyploidy 
from Herbarium Specimens 89 

Celastrus seandens 68 

Celosia argentea var. cristata 301. 
305 

Centaurium umbellatum 21 

Cephalozia leucantha 218; spini- 
gera 219; subdentata 219 

Cephaloziella arctica 223; spini- 
gera 219 

Cerastium vulgatum 65 

Ceratophyllum demersum 322 

Cetraria ciliaris 77, 82; islandica 


77, 82; Oakesiana 82; Tucker- 
mani 82 

Chaerophyllum procumbens 20 

Chamaesaracha grandiflora 113; 
nana 113; physaloides 171 

Channell. R. B. A New Species 
of Samolus (Primulaceae) from 
Mexico 1 

Chelone glabra 206 

Chenopodium album 300, 304; 
ambrosioides 300, 305; var. an- 
thenminticum 300, 305; Bonus- 
Henricus 300, 305; Boscianum 
300. 303; Botrys 300, 304; Capi- 
tatum 300, 302; carinatum 300. 
305; foliosum 300, 305; glaucum 
300. 305; graveolens 300, 305; 
humile 300, 305; hybridum var. 
gigantospermum 300, 302; inc- 
anum 300, 305; lanceolatum 65. 
300. 304; leptophyllum 300. 302. 
303; murale 300, 305; paganum 
300. 305; polvspermum 300. 305; 
rubrum 300, 304; urbicum 300, 
304 

Chromosome Numbers in the 
Genus Krameria: Evidence for 
Familial Status 101 

Chromosome Races in the Chry- 
santhemum Leucanthemum 
Complex 122 

Chrysanthemum ircutianum 124; 
lacustre 26; Leucanthemum var. 
pinnatifidum 66; leucanthemum 
122, 123, 124; var. subpinnatifi- 
dum 124 

"hrvsobalanus oblongifolius 16, 23, 
29 

Chrysoma pauciflosculosa 290 

Chrvsoma paucifloseulosa in the 
Carolinas, Striga asiatica and 289 

Chrysosplenium iowense 248, 251 

Cichorium Intvbus 66 

Cireaea quadrisuleata var. cana- 
densis 68 

Cirsium arvense 26; pitcheri 251; 
vulgare 66 

Cladonia apodocarpa 78; bacillaris 
80; f. clavata 80; f. reagens 80; 
Boryi 77; caespiticia 78; capitata 
80; carassensis 78; f. regularis 
78; chlorophaea 79; f. carpo- 
phora 79; clavulifera 81; f. nudi- 


334 Rhodor: 


caulis 81; coniocraea 79, 81; f. 
phyllostrota 79; f. stenosevpha 
79; conista 79; cornutoradiata 
79; f. radiata 79; eristatella 80; 
f. ochrocarpia 80; f. simulata 80; 
f. squamosissima 80; f. vestitu 
80; eylindrica 79, 81; f. seyphif- 
era 79; deformis 79; delicata 80; 
didyma 80; farinaceae 77; fim- 
briata 79, 81; Floerkeana 80; f. 
trachypoda 80; furcata 77, 78; 
var. palamaea 77; var. pinnata 
77; gracilis 79; Grayi 80; f. carpo- 
phora 80; f. prolifera 80; f. 
squamulosa 80; incrassata 80; 
macilenta 80; mitis 78; mitrula 
80; nemoxyna 79, 81; papillaria 
71; piedmontensis 80; f. squamu- 
losa 80; pityrea 79, 81; pleurota 
79; var. frondescens 79; pyxidata 
79; rangiferina 77; f. erispata 77; 
scabriuscula 77; squamosa 78; f. 
squamosissima 78; subcariosa 81; 
f. squamulosa 81; submitis 78; 
sylvatica 78; tenuis 77; turgida 
78; f. squamulosa 78; uncialis 
77; verticillata 79; f. aggregata 
79; f. apoctita 79; f. phylloce- 
phala 79 

Claytonia aurea 259; chrysantha 
259; flava 259; lanceolata 259; 
virginia 258, 259 

Cnidoscolus stimulosus 291 

Cochranea anchusaefolia 21 

Comandra Richardsiana 206, 298, 
303; umbellata 298. 302 

Commelina communis var. ludens 
12 

Convallaria canaliculata 292 

Convolvulus sepium 68 

Coreopsis basalis 26; drummondii 
26; verticillata 126 

Corispermum hyssopifolium 300. 
305 

Cornus alternifolia 68; florida 206 

Corylus americana 67 

Cosmos bipinnatus 26 

Crataegus erus-galli 320 

Crepis pulehra 26 

Croton capitatus var. lindheimeri 
17; engelmanni 17; lindheimeri 
17 

Cryptocolea imbricata 223 


| VoL. 60 


Cuphea balsamona 19; carthagen- 
ensis 19 

Cuscuta campestris 66 

Cycloloma atriplicifolium 14, 300. 
305 

Cynomarathrum saxatile 269 

Cyperus erythrorhizos 317; flave- 
scens var. poaeformis 206; stri- 
gosus 67 

Cytotaxonomic Study of the Genus 
Achillea in Pennsylvania 7 


Dactylis glomerata 65 

Daucus Carota 65 

Dermatocarpon aquaticum 82; 
miniatum var. complicatum 82 

Description, Distribution and 
Ecology of Three Species of 
Vaucheria Previously Unknown 
from North America 283 

Determination of Polyploidy from 
Herbarium Specimens 89 

DeWolf, G. P. Jr. Evolution of 
Flowering Plants (Review) 262 

Dicerandra densiflora 23, 29 

Dichanthium annulatum 91, 92, 94, 
Plate 1232, 95 

Digitaria filiformis 11, 30; var. vil- 
losa 208; Ischaemum 65; san- 
guinalis 65; texana 11. 30 

Diplophyllum albicans 216; apicu- 
latum 211, 228, 229, 243, 244; var. 
obtusatum 228; incurvum 216; 
obtusifolijm 228, 229, 230 

Dipsacus sylvestris 174; sylvestris 
f. albidus 175 

Distribution of Litorella americana 
in the Mid-arrowhead Region of 
Minnesota 33 

Dithvrea Griffithsi 147, 151; Wisli- 
zenii 146, Plate 1233, 147. 150. 
151. 152; var. Palmeri 151 

Dominican Republic. Thelepogon 
elegans Roth in the 73 

Dryopteris thelypteris 295; The- 
Ivpteris var. pubescens 69, 206 


Eaton. R. J. Some Interesting 
Records from Eastern Massachu- 
setts 316; The Spontaneous Flora 
of an Old House-Lot in Eastern 
Massachusetts 61 

Eaton, R. J., Bean, R. C. and Hill, 


1958] Index to Volume 60 339. 


A. F.. Twelfth Report of the 
Committee on Plant Distribu- 
tion 297 

Echinochloa pungens var. Wie- 
gandii 67 

Echinodorus parvulus 42 

Echium vulgare 43 

Ehrle, E. B, A Cytotaxonomic 
Study of the Genus Achillea in 
Pennsylvania 7 

Elaeagnus pungens 18 

Elatine triandra in New York 58 

Elatine triandra 58; f. submersa 
58. 59 

Emex spinosa 298, 305 

Epilobium coloratum 68 

Equisetum arvense 64, 69 

Eragrostis lugens 11; spectabilis 67 

Erigeron annuus 68, 72; canadensis 
68; pulchellus 68 

Eugenia jeremiensis 326 

Eupatorium fistulosum 206; per- 
foliatum 295 

Euphorbia commutata 296; corol- 
lata 206; dentata 17; hirta 18; 
Ipecacuanhae 291; supina 68 

Euphrasia disjuncta 308; micran- 
tha 307, 308; subarctica 308 

Euphrasia micrantha New to 
North America 307 

Evernia mesomorpha 76 

Evers, R. A., Further Notes on the 
Illinois Flora 142 

Evolution of a Taxonomic Charac- 
ter in Dithyrea (Cruciferae), 
The Genetic 145 

Evolution of Flowering Plants 
(Review) 262 

Experiments and Observations 
Bearing on Evolution in Oeno- 
thera 37 


Facelis apiculata 27; retusa 27 

Fagopyrum sagittatum 298, 304; 
tataricum 298, 305 

Falearia sioides 41 

Fell, E. W. New Illinois Carex 
Records 115 

Festuca elatior 65; ovina 65; rubra 
67; var. commutata 67 

Field and Herbarium Studies in 
Southern Illinois 292 

Fimbristylis autumnalis 206 


Florida, A New Species of Vicia 
(Leguminosae) in 256 

Forestiera acuminata 327; cassi- 
noides 328; porulosa 328 

Foster, R. C. Three Changed 
Authorities for Combinations 86 

Fragaria virginiana 67 

Fraxinus americana 68, 71 

Froelichia arizoniea 301, 305; flori- 
dana 14; var. campestris 208; 
gracilis 14, 41, 116 

Frullania bolanderi 211, 225, 226. 
227, 228, 229, 244, 247, 248, 252 

Fruticose and Foliose Lichens of 
Worcester County, Massachu- 
setts. The 74 

Further New Records of Myrta- 
ceae from Jamaica 323 

Further Notes on the Illinois Flora 
142 


Galinsoga ciliata 66 

Garberia fruticosa 119; heterophyl- 
la 119 

Gates, B. N., Cabomba caroliniana 
Grows in Worcester County, 
Massachusetts 321 

Gates, R. R., Experiments and 
Observations Bearing on Evolu- 
tion in Oenothera 37 

Gaura parviflora 19 

Generic Considerations Concerning 
Carphephorus, Trilisa and Litri- 
sa (Compositae) 117 

Genetic Evaluation of a Taxono- 
mic Character in Dithyrea (Cru- 
ciferae) The 145 

Geobalanus oblongifolius 16 

Geocaulon lividum 298, 302 

Geranium maculatum 68, 69; 
Robertianum 66 

Geum laciniatum var. trichocar- 
pum 67 

Glassman, S. F. and Thieret, J. W.. 
Grasses New to Illinois and the 
Chicago Region 264 

Gleason, H. A., Two New Stations 
for Carex picta 175 

Gloeocystis scopulorum 284 

Gnaphalium obtusifolium 68 

Godfrey, R. K. and Kral, R., A 
New Species of Vicia (Legumi- 
nosae) in Florida 256; Some 


336 Rhodora 


Identities in Halesia (Styraca- 
ceae) 86 

Gonolobus laevis 21 

Grasses New to Illinois and the 
Chicago Region 264 

CGutierrezia sarothrae 270 

Gyrophora Dillenii 84 


Habenaria ciliaris 206; clavellata 
206; lacera 67 

Halesia carolina 88; diptera 86; 
Plate 1231, 87, 88; var. diptera 
SNB; var. magnifolia 88; parvi- 
flora S8; tetraptera Plate 1231, 
S7. NS 

Halesia (Styracaceae), Some Iden- 
tities in 86 

Harperella fluviatilis 21 

Harris, S. K.. A New Color Form 
of Solidago sempervirens 2061; 
Cabomba caroliniana in Essex 
County. Massachusetts 116; Ne- 
lumbo Lutea in Essex County, 
Massachusetts 116; Rorippa Ses- 
siliflora in Essex County, Massa- 
chusetts 260 

Heiser, C. B. Jr, Three New 
Annual Sunflowers (Helianthus) 
from the Southwestern United 
States 272 

Helianthella parryi 267 

Helianthus agrestis 272; angusti- 
folius 126, 143; annuus 272, 274, 
275, 277; brevifolius 126; debilis 
275, 277, 279; ssp. cucumerifolius 
279; ssp. praecox 279; divaricatus 
126; grosseserratus 126; hirsutus 
var. hirsutus 294; var. trachy- 
phyllus 294; maximiliani 27; 
neglectus 274, 275, 276, 277, 279. 
282, 283; paradoxus 272, 273. 
274, 275, 277, 282; petiolaris 272. 
274, 275, 277, 279; ssp. fallax 
279. 280, 282; ssp. petiolaris 279. 
281, 282; petiolaris var. canescens 
277. 282; praetermissus 274; ra- 
dula 27; tuberosa 64, 66; x am- 
biguous 126; annuus X petiolaris 
279; H. x luxurians 126; gigan- 
teus X mollis 126; neglectus X 
annuus 277, 278; neglectus x 
debilis ssp. cucumerifolius 278; 
X debilis ssp. praecox 278; x 


| Vor. 60 


petiolaris var. canescens 278; X 
petiolaris A 278, 279; X petiol- 
aris B 278; occidentalis X grosse- 
serratus 126; petiolaris X annuus 
126 

Hehotropium amplexicaule 21 

Hepaticae VI. Phytogeographical 
Relationships of Critical Species 
in Minnesota and Adjacent 
Areas of Great Lakes, Notes on 
Nearctic 209, 243 

Herberta hutchinsiae 224; sakuraii 
210, 224; tenuis 224 

Hermann, F. J.. Euphrasia mi- 
crantha New to North America 
307 

Hesperis matronalis 16 

Heteropogon melanocarpus 11 

Hieracium aurantiacum 66; floren- 
tinum 66; pilosella var. niveum 
320 

Hilferty, F. J.. A New Bryophyte 
Flora (Review) 176 

Hill. A. F.. Eaton, R. J. and Bean. 
R. C.. Twelfth Report of the 
Committee on Plant Distribu- 
tion 297 

Hodgdon, A. R. and Pike, R., A 
New Rhododendron Station in 
Maine 328 

Holosteum umbellatum 15 

Humulus japonicus 297, 304; 
Lupulus 297, 304 

Hybrid Ragwood, Ambrosia arte- 
misiifolia X trifida, The 309 

Hydrocotyle sibthorpioides 20 

Hymenocallis occidentalis 13 

Hymenoxys richardsonii 270 

Hypericum kalmianum 251; muti- 
lum var. parviflorum 68; per- 
foratum 68; tubulosum var. 
Walteri 206 

Hypochaeris glabra, 10. 28, 29 

Hypogymnia physodes 82 


Ilex myrtifolia 18; opaca 206 

Illinois and the Chicago Region, 
Grasses New to 264 

Illinois, Buchloe dactyloides in 
259; Field and Herbarium Stud- 
ies in Southern 292; Plant Notes 
from 41 


1958] Index to Volume 60 ood 


Illinois Flora, Further Notes on 
the 142 

Impatiens capensis 68 

Ionoxalis martiana 17 

Iris brevicaulis 42 

Isoetes macrospora 316; muricata 
34; riparia 316; var. canadensis 
316 

Itea virginica 206 

Iva ciliata 28 


Jamaica, Further New Records of 
Myrtaceae from 323 

James, C. W., Generic Considera- 
tions Concerning Carphephorus. 
Trilisa and Litrisa (Compositae) 
117 

Jennings, O. E. Thelepogon ele- 
gans Roth in the Dominican 
Republie 73 

Jones, G. N., Buchloé dactyloides 
in Illinois 259 

Juglans cinerea 71; nigra 65 

Juncus diffusissimus 206; tenuis 67 

Jungermannia polaris 222; schiff- 
neri 211, 222, 223 

Jussiaea decurrens 206; leptocarpa 
19, 142, 143 


Kochia Scoparia 300, 305; Sieversi- 
ana 300, 305 

Kral, R. and Godfrey, R. K.. A 
New Species of Vicia (Legumi- 
nosae) in Florida 256 

Krameria grayi 102. 103, 105; 
lanceolata 102. 103. 105; ramo- 
sissima 102, 103. 105 


Lactuca canadensis var. latifolia 
68; var. longifolia 68 

Lakela, O., Distribution of Lit- 
torella americana in the Mid- 
arrowhead Region of Minnesota 
33 

Laportea canadensis 297, 302 

Lapsana communis 28 

Larrea tridentata 5 

Leontodon autumnalis 66; leysseri 
28 

Lepanthes Brenesii 100; cochleari- 
formi 98; turialvae 97, 98, 99, 
100 

Leptodactylon pungens 270 


Lewis, W. H.. Minor Forms of 
North American Species of Rosa 
237 

Liatris odoratissima 117, 118; pani- 
culata 118 

Lichens of Worcester County. 
Massachusetts, The Fruticose 
and Foliose 74 

Linaria canadensis 68, 206 

Lindera Benzoin 206 

Lithospermum croceum 296; caro- 
liniense 206; 296 

Litrisa carnosa 118, 119, 120 

Littorella americana 33, 34, 35, 36, 
37; uniflora 33, 36, 37 

Lobelia Dortmanna 33, 37 

Lohum perenne 65 

Long, R. W., Notes on the Distri- 
bution of Ohio Compositae: I. 
Heliantheae, Anthemideae 125 

Lophozia alpestris 253; gillmani 
213. 223, 224; grandiretis 253; 
groenlandica 253; hatcheri 253; 
heterocolpa 223; kunzeana 223. 
253; latifolia 223; lycopodioides 
253; obtusa 253; quadriloba 213. 
214, 233; schultzu 214, 224 

Lotus corniculatus 42 

Love, D. and Bernard, Fr. J-P.. 
Rumex stenophyllus in North 
America 54 

Luzula multiflora 70 

Lysimachia Nummularia 66; quad- 
rifolia 68 

Lythrum carthagenense 19 


Maguire, B. Arenaria rossii and 
Some of Its Relatives in America 
44 

Maianthemum canadense 67 

Maine, A New Rhododendron Sta- 
tion in 328 

Malus platycarpa var. hoopesii 294 

Mannia fragrans 233; pilosa 211. 
233, 234, 247, 249, 250; rupestris 
211, 243, 244, 245; sibirica 211. 
233, 234, 249, 250 

Marsupella emarginata 246; paro- 
ica 245, 246; sparsifolia 245, 246 

Massachusetts, Cabomba carolini- 
ana Grows in Worcester County, 
Massachusetts 321; Cabomba 
caroliniana in Essex County 116; 


338 Rhodora 


Nelumbo lutea in Essex County 
116; Rorippa sessiliflora in Essex 
County 260; Some Interesting 
Records from Eastern 316; The 
Fruticose and Foliose Lichens of 
Worcester County 74; The Spon- 
taneous Flora of an Old House- 
Lot in Eeastern 61 

Matricaria Chamomilla 127 

Megalodonta Beckii 322 

Mehra, K. L. and Celariar, R. P., 
Determination of Polyploidy 
from Herbarium Specimens 89 

Melica nitens 11 

Mentha alopecuroides 43 

Mertensia paniculata 226, 248, 249, 
251 

Mexico, A New Species of Samolus 
(Primulaceae) from 1 

Michigan, with the Description of 
a New Intermediate Taxon, Pe- 
rennial Ragweeds (Ambrosia) in 
177 

Microlepidozia setacea 220 

Mimulus geyeri 43; guttatus 58; 
luteus var. eupreus 86 

Minnesota and Adjacent Areas of 
the Great Lakes, Notes on 
Nearctic Hepaticae VI. Phyto- 
geographical Relationships of 
Critical Species in 209, 243 

Minnesota, Distribution of Litorel- 
la americana in the Mid-arrow- 
head Region of 33; Notes on the 
Distinction between the Broad- 
and Narrow-leaved Antennarias 
of 306 

Minor Forms of North American 
Species of Rosa 237 

Minuartia rolfii 44-46; rossii 45 

Mirabilis hirsuta 301, 305; jalapa 
14; Jalapa 301. 305; linearis 301. 
305; nyctaginea 301, 305 

Missouri, An Unusual Botanical 
Area in 205; X Carex Deamii 
in 174 

Mitrula phalloides 207 

Mohlenbrock, R. H. Jr.. Field and 
Herbarium Studies in Southern 
Illinois 289 

Mollugo verticillata 66. 301. 304 

Monachino, J.. Elatine triandra in 
New York 58 


[VoL. 60 


Monolepis Nuttalliana 300. 305 

Morley, T., Notes on the Distinc- 
tion between the Broad- and 
Narrow-leaved Antennarias of 
Minnesota 306 

Muhlenbergia frondosa 67; mexi- 
cana 67; Schreberi 67 

Mulligan, G. A. Chromosome 
Races in the Chrysanthemum 
Leucanthemum Complex 122 

Myosotis versicolor 22 

Myrcia cardiophylla 325; skeldingi 
325; uberavensis 325 

Myriophyllum tenellum 34 

Myrtaceae from Jamaica, Further 
New Records of 323 


Name Lepanthes Turialvae: A 
Source of Confusion, The 97 

Nearctic Hepaticae VI. Phyto- 
geographical Relationships of 
Critical Species in Minnesota 
and Adjacent Areas of the Great 
Lakes 209, 243 

Nelumbo lutea 116 

Nelumbo lutea in Essex County, 
Massachusetts 116 

Neoparrya lithophila 265, 269, 270; 
Mathias 265 

New Bryophyte Flora, A (Review) 
176 

New Color Form of Solidago 
sempervirens, A 261 

New Illinois Carex Records 115 

New Rhododendron Station in 
Maine A 328 

New Species of Samolus (Primu- 
laceae) from Mexico, A 1 

New Species of Vicia (Legumino- 
sae) in Florida, A 256 

New York, Elatine triandra in 58 

North America, Description, Dis- 
tribution and Ecology of Three 
Species of Vaucheria Previously 
Unknown from 283; Euphrasia 
micrantha New to 307; North of 
Mexico, A Taxonomic Study of 
the Genus Physalis in 107, 128, 
152; Rumex stenophyllus in 54; 
Some New Flora for Parts of 
(Review) 235 

North American Species of Rosa, 
Minor Forms of 237 


1958] Index to Volume 60 339 


North or South Carolina, Species 
New to the Flora of 10 

Notes on the Distinetion between 
the Broad- and Narrow-leaved 
Antennarias of Minnesota 306 

Notes on Nearctic Hepaticae VI. 
Phytogeographical Relationships 
of Critical Species in Minnesota 
and Adjacent Areas of Great 
Lakes 209 

Notes on the Distribution of Ohio 
Compositae: I. Heliantheae, An- 
themideae 125 

Nuphar variegatum 116 

Nymphaea odorata 116; tuberosa 
34 

Nymphoides peltatum 58 


Odontoschisma elongatum 219; 
macounii 212, 223 

Oenothera blandina 39; Lamarcki- 
ana 37; perangusta 37, 38, 39. 40; 
var. rubricalyx 38; Victorini 39 

Oenothera, Experiments and Ob- 
servations Bearing on Evolution 
in 37 

Ohio Compositae: I. Heliantheae. 
Anthemidaeae, Notes on the 
Distribution of 125 

(Oleaceae) Typification of the 
Genus Forestiera 327 

Onoclea sensibilis 69, 206 

Orange-Yellow-Flowered Claytonia 
virginica 258 

Ornithagalum umbellatum 65 

Orobanche uniflora 68 

Osmunda cinnamomea 69, 206; 
regalis var. spectabilis 69, 206 

Oxalis europaea f. villicaulis 68; 
martiana 17 

Oxyria digyna 298, 302 


Panicum capillare var. oceidentale 
67; commutatum 206; lanugi- 
nosum var. fasciculatum 67; var. 
implicatum 67; laxiflorum 206; 
microcarpon 206; polvanthes 
206; spretum 67 

Parietaria floridana 297. 305; pen- 
sylvanica 297. 303. 318, 320 

Parmelia Borreri 83; caperata 83; 
conspersa 83; f. insidiata 83; 
var. stenophylla 83; dubia 83; 


perforata 83; rudecta 83; saxa- 
tilis 83; subaurifera 83; sulcata 
83 

Parsonsia balsamona 19 

Parthenocissus quinquefolia 68 

Paspalum ciliatifolium var. Muh- 
lenbergii 67; plicatulum 11. 30; 
selaceum 206, 207, 208; stra- 
mineum 116 

Peltigera canina 82 

Pennsylvania. A  Cytotaxonomic 
Study of the Genus Achillea in 7 

Penstemon arkansanus 292; palli- 
dus 292 

Perennial Ragweeds (Ambrosia) 
in Michigan, with the Descrip- 
tion of a New Intermediate 
Taxon 177 

Perizoma rhomboidea 23 

Persicaria hirsuta 18 

Phalaris canariensis 12 

Phleum pratense 65 

Phlox drummondi 21; gracilis 41; 
pilosa 206 

Phyllanthus niruri 18; pentaphyl- 
lus 18, 30 

Physalis aequata 160; Alkekengi 
107, 111, 128, 133; ambigua 140; 
angulata 111. 162; var. angulata 
112. 132. 162. 163; var. lancei- 
folia 112, 132, 163; var. Linkiana 
162; var. pendula 112. 132. 163; 
angustifolia 108, 109, 129, 137; 
arenicola 109, 139; var. arenicola 
129, 139; var. ciliosa 129, 139; 
barbadensis 164; var. glabra 165; 
var. obscura 165; cardiophylla 
159; Carpenteri 128, 171; caudel- 
la 108, 112, 139. 158; ciliosa 108. 
139; comata 108. 157: crassifolia 
107. 109. 111. 112. 159. 170; var. 
cardiophylla 159; var. crassifolia 
132. 159; var. versicolor 112. 132. 
160; Elliottii 134; Fendleri 108. 
158; var. cordifolia 158; floridana 
108, 164; foetens var. neomexi- 
cana 133. 168; Francheti 133; 
fuscomaculata 134; genucaulis 
108, 160; Greenei 133, 170; he- 
deraefolia 108. 109. 111. 112, 156; 
var. comata 131. 157: var. cordi- 
folia 131. 158; var. Fendleri 111; 
var. hederaefolia 131, 156; var. 


340 Rhodora 


puberula 156; heterophylla 68. 
107, 108, 109, 110, 111, 140, 141, 
155; var. ambigua 140; var. cla- 
vipes 130, 141; var. heterophylla 
130, 140; var. umbrosa 140; var. 
villosa 130, 141; hirsuta var. in- 
tegrifolia 166; var. repandodent- 
ata 165; ixocarpa 108, 112, 113. 
128, 131, 160; lanceifolia 107, 
163; lanceolata 107, 108. 141, 155; 
var, hirta 138; var. laevigata 154; 
var. longifolia 154; var. spathu- 
lacfolia 135; latifolia 141; lati- 
physa 133, 169; Linkiana 107. 
162; lobata 110. 111, 113. 128. 
171; f. albiflora 171; longifolia 
108. 112. 154; macrophysa 108, 
112, 153; maritima 133, 134 ; mis- 
souriensis 108. 111, 133. 169. 170; 
mollis 108, 135; var. cinerascens 
136; var. parvifolia 136; monti- 
cola 108. 142; muriculata 159; 
neomexicana 108, 168, 169; nyc- 
taginea 108, 140; obscura 165; 
var. glabra 107. 165; var. viscido- 
pubescens 107, 164; Palmeri 156; 
pedunculata 170; pendula 108. 
163; pensylvanica 107, 136; var. 
cinerascens 136; peruviana 107. 
108, 130, 141; var. latifolia 141; 
polyphylla 108. 156; pruinosa 
167; pubescens 111, 112, 164. 166. 
170; var. glabra 132, 165; var. 
grisea 132, 167; var. integrifolia 
132. 166; var. pubescens 132. 164; 
pumila 108, 129, 138; var. sono- 
rie 154; rigida 108. 154; rotun- 
data 108, 157; sabeana 171; 
sinuata 108. 140; subglabrata 
108. 152; texana 153; turbinata 
164. 165; variovestita 129. 137. 
138; versicolor 108, 160; virgini- 
ana 107. 109. 111. 141. 142. 155; 
f. macrophysa 153; var. ambigua 
140; var. campaniforma 130. 
156; var. hispida 130. 154; var. 
intermedia; var. polyphylla 131, 
156; var. sonorae 111. 112, 131. 
154; var. subglabrata 111. 112. 
152, 160; f. subglabrata 130; f. 
macrophysa 130; var. texana 
131, 153; var. virginiana 111. 112. 
130, 142; viscido-pubescens 164; 


[ Vor. 60 


viscosa 107, 111, 112, 133, 136. 
137; f. Elhotti 134; f. latifolia 
134; ssp. maritima 134; ssp. 
mollis 135; ssp. viscosa 133; var. 
cinerascens 110. 129, 136. 138; 
var, Elliottii 134; f. Blliottii 129; 
f. glabra 129, 135; var. maritima 
110, 134. 137; f. maritima 129; 
f. latifolia 129; var. mollis 110. 
129, 136, 138; var. spathulaefolia 
110, 129, 135; Walteri 108. 134; 
Wrightii 108, 111, 128, 132, 161. 
171 

Physcia aipolia 83; ascendens 83; 
elaeina 83; leptalea 83; leuco- 
leiptes 83; millegrana 83; orbi- 
cularis 83; f. rubropulchra 83; 
pulverulenta 83; stellaris 84; f. 
tuberculata 84; var. rosulata 84; 
tribacia 83 

Phytogeographical Relationships 
of Critical Species in Minnesota 
and Adjacent Areas of Great 
Lakes, Notes on Nearetic Hepa- 
ticae VI. 209, 243 

Phytolacca americana 67, 301, 303 

Picris hieracioides 29 

Pike. R. and Hodgdon. A. R. A 
New Rhododendron Station in 
Maine 328 

Pilea fontana 13, 295; opaca 13. 
295; pumila 13, 67. 295. 297. 302; 
var. deamii 295 

Pinguicula vulgaris 253 

Pinus echinata 292; edulis 270; 
rigida 71; Strobus 67. 329 

Plantago aristata 24; hookeriana 
var, nuda 10, 24, 30; lanceolata 
66; major 66; patagonica 24; 
Rugeli 68; Wrightiana 24 

Plant. Notes from Illinois 41 

Pluchea petiolata 206 

Poa compressor 65; pratensis 67 

Poinsettia dentata 17 

Polygonatum biflorum 292. 293; 
canaliculatum 292; commutatum 
292, 293; pubescens 292. 293 

Polvgonella americana 206; articu- 
lata 298, 303 

Polygonum achoreum 298. 303: 
allocarpum 298. 304. 305; am- 
phibium f. terrestre 298. 305; 
var. stipulaceum 298, 302; f. 


1958] Index to Volume 60 341 


fluitans 298, 302; f. hirtuosum 
298; f. simile 298; arifolium 206; 
var. pubescens 298, 302. 303; 
aviculare 65, 298, 304; var. lit- 
torale 298, 304; var. vegetum 
298, 304; Bistorta 298, 305; 
Careyi 298, 302; cespitosum var. 
longisetum 298, 305; cilinode 298, 
302; f. erectum 298, 302; cocci- 
neum 298, 302; f. natans 298, 
302; Convolvulus 65. 298. 304; 
var. subalatum 298, 304; crista- 
tum 298, 303; cuspidatum 298. 
305; Douglasii 298, 302; dubium 
208. 305; erectum 298, 303; ex- 
sertum 299, 304; Fowler 299. 
305; glaucum 299, 304; hirsutum 
13; Hydropiper 67. 299. 302; var. 
projectum 299, 302; hydropiper- 
oides 299, 302; f. strigosum 299; 
lapathifolium 299, 302; var. pros- 
tratum 299, 305; var. salicifolium 
299, 302; minus var. subcontinu- 
um 299, 305; nepalense 299, 305; 
opelousanum 299, 303; var. ade- 
nocalyx 299, 303; orientale 299, 
304; pensylvanicum 299, 303; 
var. laevigatum 67. 299. 302; f. 
albineum 299; var. nesophilum 
299. 303; Persicaria 65, 299. 304; 
var. ruderale 299, 305; polvsta- 
chyum 299, 305; prolificum 299. 
304; punctatum 299. 302; var. 
leptostachyum 299, 302; var. par- 
vum 299, 304; puritanorum 299, 
303; ramosissimum 299, 304; f. 
atlanticum 299, 304; robustius 
299. 303; sachalinense 299, 305; 
sagittatum 299, 302; f. chloran- 
thum 299, 304 ; scabrum 299, 305; 
scandens 299, 303; setaceum var. 
interjectum 299, 303; tenue 299. 
303; viviparum 253, 299, 302 

Populus balsamea 248 

Portulaca oleracea 65 

Potentilla canadensis 67; trident- 
ata 248. 253 

Preissia quadrata 234 

Primula mistassinica 248 

Proctor, G. R.. Further Records of 
Myrtaceae from Jamaica 323 

Prunella vulgaris 66 


Prunus serotina 67; virginiana 67, 
69 

Ptilimnium fluviatile 20 

Puccinellia distans 264 

Pyrus angustifolia 293, 294; coro- 
naria 293, 294; var. lancifolia 
294; ioensis 294; lancifolia 293; 
melanocarpa 206 

Pyxine sorediata 82 


Quercus alba 67, 71; coccinea 206; 
faleata 206; imbricaria 206; leav- 
is 291; rubra 67, 71; velutina 206 

Quincula lepidota 108, 171; lobata 
171 


Radford, A. E., Ahles, H. E. and 
Bell, C. R.. Species New to the 
Flora of North or South Carolina 
10 

Ramallina pollinaria 76 

Ranunculus acris 65; arvensis var. 
tubereulatus 15; flabellaris 322; 
harvevi 292; parvulus 15; repens 
34. 37. 65; sardous 15 

Raphanus raphanistrum 16; f. al- 
bus 16 

Ratibida columnifera 116 

Reboulia hemisphaerica 230 

Rediscovery of the Genus Neo- 
parrva Mathias (Umbelliferae) 
265 

Rhamnus Frangula 61, 65 

Rheum Rhaponticum 299, 305 

Rhexia virginica 206 

Rhododendron lapponicum 251; 
maximum 328, 329 

Rhus radicans 70 

Rollins. R. C.. Bartholomew’s Cob- 
ble (Review) 32; The Genetic 
Evaluation of a Taxonomic 
Character in Dithvrea (Cruci- 
ferae) 145; Orange-Yellow 
Flowered Claytonia virginica 
258: Some New Floras for Parts 
of North America (Review) 235; 
Species Plantarum—More Than 
a Facsimile Edition (Review) 
59 

Rorippa sessiliflora in Essex Coun- 
tv. Massachusetts 260 

Rosa acicularis var. Gmelini f. alba 
242; ssp. acicularis f. alba 242; 


342 Hhodora 


ssp. Sayi f. plena 242; adeno- 
sepala 240; arkansana 241; f. 
alba 241; f. plena 241; f. setu- 
losa 241; arkansoides f. alba 241; 
blanda 240; f. carpohispida 240; 
var. hispida 240; blanda carpo- 
hispida 240; carolina 240, 242; 
f. plena 242; var. alba 240; var. 
hemisphaerica 240; var. inermis 
239; var. pimpinellifolia 240; 
var. plena 242; var. scandens 
240; var. setigera 237; carolina 
conglobata 242; carolina inermis 
239; carolina salicifolia 239; 
carolinana var. flore pleno 242; 
heliphila f. alba 241; hudsoniana 
239; hudsonia flore multiplici 
240; hudsonia salicifolia 239; 
hudsonia subcorymbosa 240; hu- 
milis var. plena 242; nitida 238; 
f. spinosa 237, 238, 239; palus- 
tris 238, 239, 240; f. alba 240; 
f. inermis 239; f. plena 240; 
var. inermis 239; parviflora 242; 
parviflora (flore multiplici) 242; 
parviflora pleno 242; pennsyl- 
vanica var. flore pleno 242; 
pennsylvanica plena 242; pratin- 
cola f. alba 241; pratincola setu- 
losa 241; suffulta f. alba 241; 
f. setulosa 241; virginiana 238; 
Woodsii f. hispida 240; blanda 
X R. pisocarpa 238; carolina X 
R. nitida 237; MacDougali x 
R. ultramontana 240; nitida X 
R. palustris 237; x R. virginiana 
237, 238, 239; Woodsiü X R. 
blanda 238 

Roubieva multifida 300. 305 

Rubus Enslenii 67; laciniatus 42; 
occidentalis 67; parviflorus 249. 
251; pensylvanicus 67; phoeni- 
colasius 42 

Rudbeckia hirta var. Brittonii 126; 
mollis 29; serotina 66; tenax 127 

Rumex Acetosa 65, 299, 304; Ace- 
tosella 65, 299. 304; alluvius 55: 
alpinus 299. 305; altissimus 299. 
304; biformis 56; brittanica 55; 
crispus 55, 56, 65, 299, 304; var. 
dentatus 56; X obtusifolius 56. 
57, 299; X orbiculatus 299; do- 
mesticus 299, 304; fenestratus 


| Vor. 60 


299. 302; hastatulatus 299, 305: 
maritimus 299, 305; var. fueginus 
300. 304; mexicanus 300. 304; 
obtusifolius 55, 56, 65, 300, 304; 
var. cristatus 56; odontocarpus 
55. 56; orbiculatus 300, 302; pal- 
lidus 300, 304; patientia 13. 
Patientia 300, 305; persicarioides 
300. 304; pulcher 55, 300. 305; 
stenophyllus 54. 55, 56, 57; tri- 
angulivalvis 300, 304; verticil- 
latus 300, 303 

Rumex stenophyllus in North 
America 54 

Ruta braveolens 17 


Sagina procumbens 67 

Sagittaria latifolia 58 

Salicornia Bigelovii 300, 304; euro- 
paea 300, 304; var. prostrata 300, 
304; var. simplex 301, 304; vir- 
ginica 301, 304 

Salsola Kali 301, 304; var. caro- 
liniana. 301, 304; var. tenuifolia 
301, 304 

Salvia pitcheri 41, 43; reflexa 41, 
43 

Samolus alvssoides 1; cinerascens 
1. 2. 4; cuneatus 1; dichondri- 
folius 2. 3. 4, 5, 6; ebracteatus 
1.2.4, 5.6 

Sassafras albidum 206 

Seapania buchii 213; curta 216; 
cuspiduligera 216, 217, 223, 224; 
degenii 220, 221, 222; var. dubia 
220. 221; gvmnostomophila 215. 
216. 223; helvetica 216; hyper- 
borea 220, 221, 222, 224; incurva 
215; microphylla 213; mucronata 
234; scandica 214, 215, 216; spitz- 
bergensis 216 

Schultes, R. E.. The Name Lepan- 
thes Turialvae: A Nource of 
Confusion 97 

Schuster. R. M.. Notes on Nearetie 
Hepaticae VI. Phytogeographi- 
cal Relationships of Critical 
Species in Minnesota and Ad- 
jacent Areas of Great Lakes 209. 
243 

Scirpus caespitosus 253; carinatus 
12; etubereulatus 12; koilolepis 
12, 30; rubricosus 206 


1958] Index to Volume 60 343 


Scleria nitida 207, 208; reticularis 
41; triglomerata 208 

Scoparia dulcis 24 

Sedum purpureum 65, 66; rosea 
219, 251 

Selaginella arenicola ssp. acantho- 
nota 291 

Sesban exaltata 17 

Sesbania exaltata 10, 17 

Setaria glauca 65 

Silphium laciniatum 127 

Sisyrinchium arenicola 319, 320; 
atlanticum 67; montanum var. 
crebrum 67 

Smilax glauca var. leurophylla 295 

Solanum americanum 68, 69; caro- 
linense 68, 69; Dulcamara 58, 66; 
rostratum 66 

Solenostoma appalachianum 245. 
246; monticola 246; sphaerocar- 
pum 253 

Solidago buckleyi 292; caesia 68; 
canadensis 62, 64, 68; houghtonii 
214; juncea 68; nemoralis 68; 
patula 206; 295; pauciflosculosa 
290; rugosa 62, 64, 68; rugosa 
var. celtidifolia 206; sempervir- 
ens f. citrea 261 

Soliva sessilis 29 

Some Identities in Halesia (Styra- 
caceae) 86 

Some Interesting Records from 
Eastern Massachusetts 316 

Some New Floras for Parts of 
North America (Review) 235 

Sonchus uliginosus 54 

South Carolina, Species New to 
the Flora of North or 10 

Species New to the Flora of North 
or South Carolina 10 

Species Plantarum—More Than a 
Facsimile Edition (Review) 59 

Spergula pentandra 15 

Sphenoclea zeylandica 24 

Spinacea glabra 301, 305; oleracea 
301, 305 

Spiranthes cernua 67 

Spirea latifolia 67 

Spontaneous Flora of an Old 
House-Lot in Eastern Massachu- 
setts, The 61 

Stachys sieboldii 23 

Stellaria media 65 


Stereocaulon coralloides 81; dac- 
tylophyllum 81; evolutoides 81; 
pileatum 81 

Steyermark, J. A., An Albino Form 
of Dipsacus sylvestris 174; An 
Unusual Botanical Area in Mis- 
souri 205; X Carex Deamii in 
Missouri 174 

Stillingia aquatica 18 

Striga asiatica 298, 290 

Striga asiatica and Chrysoma 
pauciflosculosa in the Carolinas 
289 

(Styraceae), Some Identities in 
Halesia 86 

Subularia aquatica 34 

Suaeda americana 301, 304; line- 
aris 301, 304; maritima 301, 304; 
Richii 301, 304 

Syntherisma filiforme 11 


Talinum calycinum 292 

Tanacetum vulgare 66 

Taraxacum erythrospermum 66; 
officinale 66 

Taxodium ascendens 18 

Taxonomic Study of the Genus 
Physalis in North America North 
of Mexico, A 107, 128, 152 

Terrell, E. E., Striga asiatica and 
Chrysoma pauciflosculosa in the 
Carolinas 289 

Tetragonia expansa 14, 301, 305 

Thelepogon elegans 73 

Thelepogon elegans Roth in the 
Dominican Republic 73 

Thieret, J. W. and Glassman, S. F., 
Grasses New to Illinois and the 
Chicago Region 264 

Thlaspi perfoliatum 16 

Three New Annual Sunflowers 
(Helianthus) from the South- 
western United States 272 

Thuja occidentalis 248 

Tovara virginiana 300, 303 

Tradescantia bracteata 42 

Tragopogon dubius 29 

Trichomanes boschianum 143, 144 

Trichostema dichotomum 43; se- 
taceum 207, 208 

Trifolium agrarium 65; pratense 
65; repens 65 


LM 


344 Rhodora 


Three Changed Authorities for 
Combinations 86 

Trilisa carnosa 120; odoratissima 
118, 119, 121; paniculata 119, 121 

Trillium recurvatum f. Shayi 42 

Trisetum pensylvanicum 207 

Triteleia uniflora 13 

Tritomaria exsectiformis 234; 
quinquedentata 253; scitula 253 

Tsuga canadensis 243, 244, 329 

Turner, B. L., Chromosome Num- 
ber in the Genus Krameria: 
Evidence for Familial Status 101 

Twelfth Report of the Committee 
on Plant Distribution 297 

Two New Stations for Carex picta 
175 

Type of Vaucheria compacta var. 
koksoakensis, The 329 

Typification of the Genus Fores- 
tiera (Olaceae) 327 

Typha latifolia 207; f. ambigua 207 


Ulmus americana 67, 71 

Umbilicaria Dillenii 84; mammu- 
lata 84; Muhlenbergii 84; pa- 
pulosa 84 

Unusual Botanical Area in Mis- 
souri, An 205 

Urocoecus foslieanus 284 

Urtica chamaedryoides 298, 305; 
dioica 298, 304; gracilis 298, 302; 
procera 298, 302; urens 298, 304; 
viridis 298, 302 

Usnea florida 81; hirta 81 


Vaccinium angustifolium var. lae- 
vifolium 68; corymbosum var. 
fuscatum 86 

Vaucheria compacta 283, 285, 286, 
287, 288; var. koksoakensis 285, 
286, 287, 288, 329: dichotoma 
287; f. marina 287; intermedia 
284; minuta 284; sphaerospora 
283, 284, 285, 287, 288; submarina 
285, 287, 288 

Verbascum Thapsus 66; virgatum 
24 

Verbena brasiliensis 22; rigida 22; 
urticifolia 68; Verbena x engel- 


8 


[ Vor. 60 


manni 22; V. hastata X V. urti- 
eifolia 22; Verbena x hybrida 
22 

Verbesina encelioides 29 

Vernonia missourica 320 

Veronica serpyllifolia 66 

Vicia acutifolia 256, 257, 258; 
Craeca 65; floridana 256, 257, 
258; ocalensis 256, 257, 258 

Viola cucullata f. albiflora 68; pa- 
pilionacea 70 

Vitis aestivalis 61, 68, 72 


Wagner. W. J. Jr, The Hybrid 
Ragweed, Ambrosia  artemisii- 
folia X trifida 309 

Wagner, W. H. Jr. and Beals. T. 
F., Perennial Ragweeds (Am- 
brosia) in Michigan, with the 
Description of a New Intermedi- 
ate Taxon 177 

Wahlenbergia gracilis 25 

Warea cuneifolia 16 

Waterfall, U. T, A Taxonomic 
Studv of the Genus Physalis in 
North America North of Mexico 
107, 128, 152 

Weber, W. A., Rediscovery of the 
Genus Neoparrva Mathias (Um- 
belliferae) 265 

Wilee, R. T. and Blum, J. L. 
Description, Distribution and 
Ecology of Three Species of 
Vaucheria Previously Unknown 
from North America 283; The 
Type of Vaucheria compacta 
var. koksoakensis 329 

Wilson, K. A., Typification of the 
Genus Forestiera (Oleaceae) 
327 

Winterringer, G. S, Plant Notes 
from Illinois 41 

Wolffia columbiana 318; punctata 
318, 319 

Woodwardia areolata 206 


Xanthoria candelaria &1 
Ximenesia encelioides 29 


Zizania aquatica 42